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Full text of "Project Coral Fish Looks at Palau"

\3 Biodiversity 
^Heritage 
^^Library 

http://www.biodiversitylibrary.org/ 



Annual report of the Board of Regents of the Smithsonian Institution. 

Washington :Smithsonian Institution, -1965. 
http://www.biodiversitylibrary.org/bibliography/7964 



1 956: http://www.biodiversitylibrary.org/item/33453 

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Project Coral Fish Looks at Palau 



By Frederick M, Bayer 

United States National Museum 
Smithsonian Institution 

and 

Robert R. Harry-Rofen 

The George Vanderbilt Foundation 



[With 20 plates] 



We who live on continents can rarely appreciate the vastness of 
the world's oceans. Those of us who may be prompted by business or 
pleasure to traverse them learn of their two-dimensional magnitude, 
but there are a few of us who are privileged to investigate the secrets 
of the seas first-hand, by living among them. We look upon the seas 
in their role as an environment and seek to unravel the interwoven 
facts of life within them. One of the first facts we learn is the com- 
plexity of their many-faceted wonders, and we consider ourselves 
fortunate when we are able, as it w^ere, to polish a few of these facets 
so as to see more clearly into them. 

Of all the seas, the one with the greatest area, greatest depth, and 
most to tell us is that restless giant, the Pacific. Even before we begin 
to study it we must concede, if not defeat, at best a draw, for it is prac- 
tically axiomatic that one solution leads to another problem. When 
our words are as antiquated as those of the Renaissance pioneers Belon 
and Kondelet and Marsigli now seem to us, people will still be learning 
new things about this watery one-third of our planet. 

When we look upon the Pacific, or any ocean for that matter, as an 
environment, the central problem deals with the physical and chemical 
properties of the fluid medium that make it adequate to support life. 
This is a vast field of investigation to which many people in many 
laboratories are devoting tireless efforts. We, however, as biologists, 
can devote time to such problems only when they have direct and im- 
mediate bearing upon some question involving the organisms with 
which we are concerned, and even then we must rely upon specialists 
in those restricted fields for most of the information we need* The 

481 



482 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1056 

next problem is a qualitative one regarding the population of the sea: 
what are these creatures that populate the waters? This is a basic 
problem in marine biology ; upon it depend the solutions to problems 
of economic, or ecologic, or purely biologic interest- We must know 
what organisms we are working with before we can determine how 
they live together in communities, how they depend upon one another, 
and how they affect us. 

Creatures of the sea do, in fact, affect the affairs of man in many 
ways. Many of them have served us for food since the beginning of 
mankind. When we build structures in the sea for our own purposes, 
certain of the animals and plants whose domain we have invaded use 
those structures to their own ends and thus either destroy what man 
has made or so befoul and beclog it as to render it worthless. When 
we sail in tropic waters, other marine life— corals and algae — has been 
there long before us and raised up an edifice that passively awaits the 
unwary navigator and his fragile keel. If we are thrown into the sea, 
or when we voluntarily venture into it, still others may unintentionally 
do us bodily damage or even deliberately seek us out as a meal. Most 
"dangerous" of all, to marine biologists anyway, are those that have 
such bizarre or complicated ways of life that they entice us to devote 
most of our lives to learning of them and solving their riddles. 

Since the close of World AVar II, interest in the Pacific Ocean has 
been increasing steadily, A number of expeditions were sent out to 
study the tropical Pacific, among which were those of the Pacific 
Science Board (National Academy of Sciences-National Research 
Council) and the George Vanderbilt Foundation. Among the 
expeditions of the Pacific Science Board were those comprising the 
5-year Coral Atoll Program, in some of which the present authors 
participated, 

The expeditions of the Coral Atoll Program did much to broaden 
our knowledge of life on the coral atolls of the Marshalls, Gilberts, 
Carolines, and Tuamotus. But field team studies came to a close 
before the most interesting part of the Pacific could be studied : the 
western rim, the f aunal gateway to that vast coral world that reaches 
to Hawaii, the Galapagos, and even our own western shores. Moving 
eastward from the Malay Archipelago and its wonderfully rich fauna, 
we find no depletion through the Philippines, but what of the western- 
most islands of Micronesia? They are a scant 600 miles east of the 
Philippine Islands (see map, fig. 1) , no journey at all for sea creatures 
with free-swimming young stages. However, between the Philippines 
and the Palau Archipelago lies one of the greatest deeps in all the 
seas. How many of the East Indian species have been able to span 
this deep? Even the submarine ridge upon which the Palaus are 
situated, extending northeastward from the Moluccas, is covered by 



PALAU — BAYER AND HARRY-ROFEN 



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484 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

waters too deep to serve as a shallow-water passageway for littoral 
forms, Are the Palaus now so isolated that special endemic species, 
found nowhere else, have evolved from their East Indian ancestors? 
How do the reefs, the habitat of most tropical shallow-water animals, 
as we see them in Palau, differ from those elsewhere in Micronesia, 
and from those in the East Indies ? Such were the questions we hoped 
to answer when we, as part of a 4 -man team, set out for the Palau 
Islands late in June 1955, 

PROJECT CORAL FISH 

Together, we formed Project Coral Fish, a continuing field program 
devoted to the study of the marine biology of the high islands and 
atolls in the Trust Territory of the Pacific Islands. This program was 
initiated in 1954 by the George Vanderbilt Foundation at Stanford 
University, with the cooperation and support of the Pacific Science 
Board (National Academy of Sciences-National Research Council), 
the Office of Naval Research, the United States Department of the 
Navy, the United States Trust Territory of the Pacific Islands, and 
the Smithsonian Institution- Aside from the present writers, the team 
included H. Adair Fehlmann, assistant curator of the George Vander- 
bilt Foundation collections at Stanford University, and Sterling H. 
Pierce, technical assistant, We hoped to gather data and specimens 
of all kinds to give us an approximate answer to the question : What 
lives in the waters of Palau? With this basic information we would 
be in a better position to answer such practical questions as: What 
kind of marine life can be exploited for food? How can their supply 
be conserved? What other marine products of economic importance 
could the Palauan people develop successfully ? What are the dangers 
in fishing the reefs, and how can they be avoided? 

As a byproduct of our basic task of finding out what animals popu- 
late the reefs of Palau, we studied the communities living in various 
kinds of habitats, especially in Iwayama Bay, and the strange associa- 
tions that develop between different kinds of animals. 

After many months of preparation, during which supplies and 
equipment were assembled and shipped to the western Pacific, the four 
team members assembled on June 22, 1955, at the George Vanderbilt 
Foundation headquarters on the Stanford University campus, and 
final plans for the trip to Palau were made. Under orders from the 
Chief of Naval Operations, the following day we left Moffett Field, 
Calif., aboard a military transport plane bound for Guam, 

Two days and more than 6,000 miles later, we arrived in Guam, 
where we learned that the expedition equipment had all been for- 
warded to the Palaus on schedule. D, PL Nucker, the Acting High 
Commissioner of the U. S. Trust Territory of the Pacific Islands, with 



PALAU — BAYER AND HARRY- ROFEN 485 

whom arrangements for the expedition had been made months pre- 
viously, issued the permits necessary for our travel and field work in 
the islands. All was in readiness. We were in the Tropics again, and 
our destination was almost in sight. 

It was an impatient group that awaited the next weekly flight of 
the Transocean Airliner to Koror, but at last Thursday came and 
we were at the air terminal in the Naval Air Station, Agana, early 
that morning. A few moments after Miss Thelma Gorman, of Trust 
Territory Headquarters, had bustled us and the other passengers 
aboard the Albatross amphibian and breathed a sigh of relief, we were 

airborne. 

The islands of Palau lie some 800 miles southwest of Guam, a flight 
of about six hours, including stops at Ulithi Atoll and Yap. At 
Ulithi, we touched down on the airstrip to discharge passengers, 
where two years previously we ourselves had landed on our way to 
Ifaluk, a tiny jewel in the sea that captured our hearts as could no 
other spot. But that is another story, and we were back in the air 
before we could reminisce about it. At Yap, the lagoon is the air- 
strip and we made a water landing to discharge passengers, cargo> 
and mail. Again, after a pause of only a few minutes, we were taking 
off on the last leg of our journey halfway around the world. The 
next land we would see would be Palau — first the great, hilly island 
Babelthuap, then Koror with its settlement and the broad harbor on 
which we would land. Reaching up toward us from the sea below, 
were the jagged ridges of the islands we would come to know so well 
(pi. 1, fig. 1)* Amid sheets of spray we settled in the green water of 
the lagoon and taxied up to the ramp that led the plane out of the 
water. At last we were in Palau, 

THE PALAUS 

Although the Palau Islands were discovered in 1710 by Francisco 
de Padilla and his pilot, Joseph Somera, on the galleon La Santisima 
Trinidad (and may have been sighted earlier, perhaps by Diego de 
Rocha in 1525-26 or Lope Martin in 1566), the first Europeans to 
publish an extensive account of their visit were the crew of the 
British East India Company's Antelope in command of Capt, Henry 
Wilson. The keel of the luckless Antelope struck the coral rocks 
of the barrier reefs near Aulong Island on the morning of August 10, 
1783. Only one man was lost in the disaster, and the remainder of the 
crew escaped safely to set up camp on Aulong, The castaways salvaged 
every usable item from the wreck and, although the Antelope was a 
total loss, were able to build another vessel large and seaworthy enough 
to take them all safely to Macao, This vessel, the Oroolong, brought 
700 Spanish dollars (about equivalent to the XL S. dollar of the 



486 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 





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Figure 2.— The Patau Islands. Adapted from U. S. H. O. chart 6073. 



SMITHSONIAN REPORT, T956. --BAYER AND HARRY-ROFEN 



PLATE 1 







1. Koror from the air, with hvayarna Bay and the limestone Islands in the distance, 
abrupt headland on the horizon at right is Ngaremediu Point on Urukthapel. 



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2, Looking east through Ngasaksao Pass, the eastern entrance to Iwayama Bay. To the 
left is Ngalap region of Koror, and to the right Kwannon (Ugeliungs) Island, with conical 
Ngaraglbukl (Xgergelbakl) Rock in the pass. 






SMITHSONIAN REPORT. 1956, — BAYER AND HARRY-ROFEN 



PLATE 2 







I. The tenore in Letnigol Passage, the narrow western entrance to Iwayama Ray. This 
boat was used by field parlies to reach the more remote localities investigated in the 

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2. A building that was once part of the Japanese meterological station on Koror now accom- 
modates biological activities in the Palau District of the IL S* Trust Territory* The 
ground-floor laboratory to the left of the main entrance is occupied by the George Vander- 
bilt Foundation, and the apartment above (now remodeled) provides living quarters for 
members of its field parties, 



SMITHSONIAN REPORT, T956- BAYER AND HARRY-ROFEN 



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PLATE 8 




1. One of the authors preparing to descend in a self-contained underwater breathing 

apparatus (aqualung)* 




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clinefish. The small urnlike objects to the left of it are colonies of an ascidian, Didemnum 



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SMITHSONIAN REPORT, 1956— BAYER AND HARRY-ROFEN 



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PLATE 10 




1. Shallow reefs in Iwayama Bay are in some places dominated by the large poritid coral 
Goniopora. The stubby branches of its skeleton are obscured by the exceptionally large 
polyps, which remain extended even in daylight. I nlike most anihozoan corals, it can 
sting, thus making collecting near it very unpleasant* 




2. This species of mushroom coral, Fungia actiniformis palawensis, has long, creamy-yellow 
tentacles that completely hide its stony skeleton which is nearly the size of a saucer. It 
is a solitary polyp that never forms a colonial skeleton as most other reef corals do. A 
small specimen of a wrasse, Ckeilinuij that is common in enclosed bays like Iwayama, can 
be seen at the top of the picture. 



PALAU — BAYER AND HAERY-ROFEN 487 

period) when sold in Macao, and stands as a tribute to British forti- 
tude and resourcefulness in the face of adversity. 

The English, although at first apprehensive, found no difficulty 
in establishing friendly relations with the Palauans. Even though 
they "had not on board philosophers, botanists, draughtsmen, or gentle- 
men experienced in such scientific pursuits as might enable them to 
examine with judgment objects which presented themselves, or trace 
nature through all her labyrinths," they nevertheless gave us a fasci- 
nating account of the islands and people and their experiences among 



Palau and the Palauans today present a far different impression 
from that described by Wilson and his men in 1783, Although the na- 
tives had complete power over the English, they did not take advantage 
of their superior position, but instead did everything possible to help 
their hapless visitors. Admittedly, the Palauans were awed by fire- 
arms, and the English assisted Abba Thulle (Ebadul or Ibedul) the 
chief of Koror, in some of his interisland campaigns. Good relations 
were thereby strengthened, but it probably was not guns alone that 
made the Palauans friendly. We ourselves have been residents of 
islands where the people have had little real contact with civilization 
as we know it, and found that they remain much as Wilson pictured 
the Palauans, with a high regard for honesty and respect for their fel- 
lows. Now the Palauan people have become sophisticated and mun- 
dane after 20 years of German administration and another 26 under 
the Japanese- Missionaries long ago insisted that they give up their 
native way of dress (or undress) in favor of less practical European- 
style clothing. Their airy, thatched houses and abais, or men's houses 
(ph 3, fig. 1), decorated with colorful murals depicting historical and 
mythological events or droll folk tales, have given way to quonset huts 
and quasi -Japanese frame structures. Their elegant outrigger canoes 
are becoming a rarity, replaced by dirty little diesel boats like the 
My Flower — anything but fiowerlike — that plies between Peleliu and 
Koror, 

The Palau Archipelago in the western Caroline Islands (fig. 2) ex- 
tends northeastward from about latitude 6° 53' N, to 8°06' N,, or 
over 70 miles, at a longitude of about 134° 29' E. The main islands, 
dominated by Babelthuap, lie between latitudes 7° and 7°45' N. Ex- 
cept for Kayangel, the northern part of the islands is volcanic in 
origin, Babelthuap is about 25 miles long and 8 miles wide, covered 
with rugged hills and dense jungle, rolling grasslands, sparkling 
streams and dashing water falls, (pi. 12, fig* 1) , and fringed with man- 
grove. Twenty miles to the north of it is Kayangel, a true atoll like 
the islands far to the east, and to the south of it is the maze of lime- 
stone ridges and conical islets (ph 1) that forms one of the most re- 

412576—57 32 



488 



ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 




Sketch of old abais, based on several published photographs* 

markable features of Palau. Here and there on the edge of the reefs 
there are islets of wave-tossed coral debris and sand, much like the 
atoll islands. Together, the Palaus offer the most diversified collec- 
tion of marine habitats to be found anywhere in Micronesia. 

The Japanese recognized the unusual scientific opportunities pre- 
sented by the Palaus and, during their administration of the South 
Seas Islands (Nanyo Gunto) , established at Koror a tropical biological 
station devoted to the study of coral reefs, under the directorship of 
Dr, Shinkishi Hatai. At that time, Koror was the capital of the 
Japanese mandated islands of the South Seas and was a thriving city. 
The Palao Tropical Biological Station is no more, but we hope to build 
upon its accomplishments and contribute further to a more thorough 
understanding of the fauna of Palau. 

ACTIVITIES IN THE FIELD 

Our first big job on Koror was to unpack and organize our field 
equipment. The territorial entomologist, Kobert P, Owen, had gen- 
erously provided us with laboratory space in the entomological labora- 
tory (pL 2, fig. 2), one of the few bullet-scarred Japanese buildings 
still usable* A rebuilt second story over the laboratory served as our 
quarters until a nearby house could be made ready for occupancy. For 
days we moved boxes and steel drums full of supplies from the district 
warehouse to the laboratory grounds, and the unpacking went on far 
into our first few nights in the islands. 



PALAU — BAYER AND HARRT-ROFEN * 489 

One morning while we were busy setting up the laboratory with 
the help of Bob Owen's Palauan assistants, we became aware oi a 
scowling stranger who at first hesitated around the periphery of ac- 
tivity, and then picked up a hammer and began energetically opening 
boxes. This was Rikrik (pi, 18, fig* 1), We asked the others about 
him and learned that he was a willing and able worker, knowledgeable 
in English, Palauan geography, and fishing. We had learned from 
experience that such a man is indispensable to an expedition, so we 
hired him on the spot. Rikrilrs scowl broke into a broad but tempo- 
rary grin and we had gained a true friend- Later, we added to our 
staff another Palauan named Sumang, who had a remarkable knowl- 
edge of Palauan natural history- He could speak both English and 
Japanese, was a village chief or "Ya'at," and knew practically every- 
body from Angaur to Kay angel. Amiable Sumang Y. was a valuable 
public-relations department whose good offices were a great advantage, 
particularly during the long overland trips on Babelthuap, and his 
memory of Palauan geographic names gave valuable documentation 
for our collection records. 

The Palauan people use a different approach from ours to naming 
the various parts of their homeland. They often do not give names 
to islands as a whole, whereas groups of islands or localized regions on 
islands may have special names. Rivers and streams may have as 
many as three names — one for the part near the mouth, another for the 
headwaters, and a third for the parts between. The imposition of our 
own practice of giving a single name to geographical features upon 
the Palauan system has led to either a part taking the name of the 
whole, or the whole taking the name of one of its parts. Examples of 
the latter kind are Koror, which is the name of a village that we apply 
to an entire island, and Eil Malk, the name of a cape which we use 
for the island of which it is a part. The situation is complicated by 
the circumstance that we take many of our spellings of Palauan place 
names from Japanese maps, which expressed them in phonetic 
katakana characters. The English transliterations from the Japanese 
spellings usually bear little, if any, resemblance to actual pronuncia- 
tion, but they appear almost universally on American maps so we are 
obliged to employ them in this account. Thus, the name "Ankosu" 
as we use it is correctly spelled "Nghus," and "Geruherugairu" should 
be "Ngaregelngael," A complete list of the place names we will men- 
tion in these pages, giving the correct (and any common alternate) 
spelling, may be found on p. 507. 

Actual field work could not be started until our 18-foot fiberglass 
boat was put into commission. Sterling H. Pierce, our engineer and 
electronics technician, installed wiring, instruments, and cabin con- 
trols for the powerful outboard motor. In due time, the final coat 



490 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

of paint was applied, the name Lenore and the George Vanderbilt 
Foundation insigne inscribed on her bows, and the boat was ready 
for launching. She was both speedy and seaworthy, and in her we 
could make trips to outlying islands and return to the laboratory the 
same day, a distinct advantage when perishable specimens must be 
promptly preserved. 

When the Lenore was fully loaded for a day's work, there was 
scarcely room left for us. Nets, containers large and small, inflatable 
floats for receiving specimens as we took them from the water, spray 
apparatus for distributing the poison we used to stupefy fishes and 
other active specimens, tools for digging and breaking up coral, pre- 
servative, cameras and, often, bulky diving apparatus loaded the boat 
to capacity. 

The self-contained diving apparatus was useful not only for col- 
lecting in water too deep for free diving, but also in shallow-water 
areas where we wanted to see the exact situation under which certain 
animals were living, or to observe their behavior at length. When- 
ever possible, both the habitat and the inhabitants at collecting sta- 
tions were photographed in detail, both in color and in black-and- 
white, using reflex cameras in waterproof casings (pi. 7, fig. 2) . These 
excellent cameras greatly minimized waste of film by enabling us to 
watch moving specimens until they were in range and focus. 

Because one of our chief aims was to get as complete a biological 
sample as possible, every available means of collecting specimens was 
employed, from hand capture and hook-and-line to explosive charges. 
Although different situations required different techniques of sam- 
pling that had to be carefully decided upon before attempting to col- 
lect, the most generally useful method of obtaining active specimens 
was by the use of the vegetable poison rotenone. In liquid form this 
is extremely potent, so it must be diluted with water in a spray pump 
and distributed over the area to be collected. Fishes, crustaceans, 
cephalopods, and certain other types of animals are suffocated by it 
and are soon made helpless. When we used this technique, we needed 
every available hand to collect specimens before they were swept away 
by the current or eaten by larger fishes not affected by the poison. 
Often we enlisted the aid of youthful spectators, who are character- 
istically good collectors, and they would scurry about in response to 
Sumang Y.'s commands, enjoying all the bustle of excitement. 

IWAYAMA BAY 

Our program of fauna! sampling took us the length of Palau, from 
Arekolong Peninsula at the northern end of Babelthuap to Peleliu, 
but one of the most fascinating and complex areas in the islands was 
virtually in our own front yard. The Japanese scientists of the 



PALAU — BAYER AND HARRY-ROFEN 491 

Palao Tropical Biological Station were attracted by it 20 years ago 
and made a survey of it that was reported in the first volume of their 
journal (Abe, 1937) . We became greatly interested in that study and 
had decided, even before reaching Palau, to resurvey the same area to 
see what had happened to the different habitats since they were origi- 
nally studied. This area is an island-studded lagoon, partly enclosed 
by Koror Island (pi. 4), which the Japanese called Iwayama- wan, or 
"Rocky-mountain Bay." The Japanese name is generally accepted 
now, because the nearly forgotten original Palauan name for it is a 
matter of debate (some maps use Kramer's coined name for it, "Songel 
a Lise," which, although utilizing Palauan words, is a European inven- 
tion). We therefore use Iwayama Bay as a neo-Palauan name. 

Iwayama Bay is a roughly circular body of water about V/ 2 nautical 
miles in diameter, enclosed by Koror on the north and Auluptagel on 
the south (see map, fig. 3). Its west entrance is a long, narrow pass 
called Lebugol Channel, and its east entrance a wider, coral- and sand- 
choked passage called Ngasaksao Pass. The western arm of Koror 
is volcanic land with a wide, muddy mangrove shore; its eastern arm 
is limestone, like Auluptagel and the 40 small islands in the Bay, with 
a fringe of corals. The Palauan names for most of the islands are all 
but forgotten (Suxnang Y. succeeded in tracking down most of them 
by lengthy conferences with the prominent patriarchs of Koror) so, 
to simplify matters, the Japanese students assigned each island a 
number, which they actually emblazoned on them in white paint. 
These roman numerals are still legible on some of the islands, and we 
also found that system more convenient to use than names, either 
Palauan or Japanese, N. Abe and his colleagues further divided the 
Bay into "divisions" bearing letter designations (see fig. 3), in most 
of which they studied transects from island shore to reef margin. In 
the process of collecting, we revisited each of the transects in the bay 
and studied the 16 divisions, wherever possible taking photographs 
of the coral growth, animal communities, and general habitat. 

Because of the many islands and narrow passes, tidal currents are 
swift at many points in Iwayama Bay. The Islands, which are very 
close together, rise up almost vertically from the bay floor and create 
very narrow, deep waterways many of which are 100 feet or more in 
depth. The islands are deeply undercut at the high- tide line (pi. 6, 
fig. 2) — as much as 5 to 10 feet — forming deep "notches' 3 above a sub- 
marine shelf of variable width. In favorable localities where the cur- 
rent is strong, as on the north shore of island 29, coral growth on the 
shelf and vertical submarine cliff is exceptionally luxuriant. Here, 
one could stand among flourishing corals and look either directly 
overhead into dense jungle vegetation or straight down the coral preci- 
pice into a hundred feet of deep blue water. Occasionally we saw the 



492 



ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 




PALAU — BAYER AND HARRT-ROFEN 493 

shadowy shapes of giant blue parrotfishes, perhaps 6 feet long, rise 
from the depth to which they had fled at our coming, or a shark patrol- 
ling his accustomed beat, or the faint silvery glint of sunlight on a 
distant school of swift predators, perhaps some kind of jack (Car- 
angidae) , near the limit of our vision. 

On the submarine shelf of these undercut shorelines, the coral 
variety was great, with multicolored Peciinm lactuca^ commonly called 
lettuce coral (pL 9, fig, 1), green, orange, or brick-red Lobophyllia^ 
and a few others dominating the shallower areas. The less colorful 
Plerogym, with bubblelike tentacles an inch across, the closely re- 
lated Physogyra^ and long, wiry antipatharians (black corals) were 
usually found on the vertical face of the cliffs. On the shadiest slopes 
we often met with handsome specimens of PalauphyUia^ a subgenus 
of corals named in honor of their homeland. 

The bright colors of Palauphyttia, like those of most other stony- 
reef corals, are located not in the coral skeleton but in the soft tissues 
of the polyps themselves, and are due in part to the presence of minute 
unicellular algae living in the cells of the endoderm. These remark- 
able algae, called zooxanthellae, actually serve the coral polyps in the 
capacity of excretory organs by taking up from the animal tissues such 
waste products as they can use in their own life processes. They have 
never been found living free of corals, and have never been artificially 
cultured. Their only reproductive process seems to be simple division 
and they are passed on from generation to generation of corals 
through the eggs, which become infected before leaving the parent. 

Several species of fishes were found only in such situations in Iwa- 
yama Bay, Among these were the brilliant gold- anti-black striped 
butterflyfish (Chaetodon octofasciatus Bleeker), wrasses of the genus 
Cheilinus (pi. 10, fig. 2), and several kinds of cardinal fishes belonging 
to the genus Apogon. Many species of highly colored fishes that reach 
a length of no more than one or two inches balance like jeweled 
spangles among coral branches that rival them in beauty, or conceal 
their splendor in holes in the coral cliffs. Ever present are the preda- 
tors that seek out these defenseless inhabitants of the coral slopes. 
Some, like trumpetfish, have long snouts with which they relentlessly 
explore all holes and crevices in search of prey; others, such as the 
turkeyfish, have wide jaws and gaping mouth that enable them to en- 
gulf their prey in one quick gulp, in much the same way that a 
vacuum-cleaner may inhale a feather* Solitary sharks were patrolling 
the bottom of the deep waterways, but their presence was of most 
importance to the inhabitants of the deeper waters for the sharks sel- 
dom came to the surface. 

Some of the islands in Iw T ayama Bay have protected little bay lets 
in which the coral growth may consist largely of Gordopora^ a thick- 



494 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1966 

branched stony coral with huge, protruding polyps (pi. 10, fig. 1). 
We soon learned to expect trouble when we collected these secluded 
spots, for the unusually virulent nematocysts, or stinging capsules, of 
the Goniopora polyps were easily dislodged by the currents we pro- 
duced by swimming, and stung us so badly that we sometimes were 
obliged to leave the water. 

Much more troublesome than Goniopora^ however, was an arbor- 
escent, colonial scy phis torn a (called Stephanoscyphm) of the medusa 
Nausithoe, which also released its nematocysts into the water upon 
agitation by strong currents. The stings produced by this animal left 
angry red welts that itched for days afterward, and even caused swell- 
ing of adjacent lymphatic glands. It seemed probable to us that a 
dense growth of Stephanoscyphm could liberate enough nematocysts 
into the water to inflict serious stings upon unsuspecting swimmers. 
Fortunately, this coelenterate seems to be common only locally in 
Palau, where we found it in only two localities. 

In spite of seemingly ideal surroundings in most parts of Iwayama 
Bay, dead corals could be found at almost any location, suggesting 
that conditions are not always so favorable. A day or so of heavy 
rainfall will dump tons of fresh water not only into the bay but also 
upon the islands, from which it cascades down to the Bay, carrying 
with it great quantities of silt and forest debris. Salinity must be 
much reduced, especially near the surface, for many hours, if not 
days; the normal water temperature of about 85° F, may be lowered 
by 5° or more ; suspended matter beclouds the water and the more deli- 
cate corals may be smothered* But, after a few clear days and several 
tidal cycles, the water clears and the survivors continue their struggle 
for existence. 

The northwest corner of the Bay, forming divisions M, N, and O, 
is bathed by good tidal flow, but silt from the nearby mangrove 
shore discourages coral growth and the reef has a sickly appearance 
that belies the large number of species that comprise it. One of the 
resurveyed transects crosses the reef-flat at the south end of island 15, 
and we studied it during several low tides. Here we collected dead 
coral heads with cylindrical black sponges growing on them, which 
we soon found were only the external portions of a boring sponge 
that had excavated great hollow caverns in the coral boulders. Some 
coral heads were thus reduced to hollow shells, and we have yet to 
learn what becomes of the sponge after it has completely "eaten itself 
out of house and home" — whether it then assumes a massive, free- 
living form as do some other boring sponges, or simply dies of expo- 
sure. Whatever its fate after the destruction of the rock in which 
it lives, it is certainly an active reef -destroying agent At the same 
locality we found another rock-boring sponge that attacked not only 



PALAU — BAYER AND HARRY-ROFEN 



495 



dead corals but also the limestone floor of the tide-level notches. 
This sponge is almost certain to prove to be one of the important bio- 
logical agents contributing to the formation of undercut shorelines. 
The swift tidal currents that breath life and variety into the reefs 
do not reach the upper part of division K in the northeast corner of 
Iwayama Bay, which is a quiet backwater. An abundant growth of 
coral is nevertheless present, but it lacks variety, Fingery masses 
of Porites and great, white, papery chalices of Montipora (pi. 11, 
fig. 2) flourish everywhere, but few others can be seen. Even the 
fishes are fewer in kind and smaller in size. Small dragonets (Cal- 
lionymidae) dart about in sandy patches or seek refuge among the 
coral branches. One of these dragonets (Synchiropus splendidus) is 




Figure 4. — The splendid dragonet, Synchiropus splendidus t slightly larger than natural size. 

probably the gaudiest fish we found during the summer — colored blue, 
green, and red in an intricate design of spots and bands (see fig- 4)* 
A few species of damselfishes {Abudefduf) were abundant, each fish 
with its own territory of coral and water. Each individual vigorously 
defends its own particular home, bullying intruders with threatening 
advances, cheeks distended, clicking and grunting indignantly. 
Hovering over the branches of Porites we could always find little 
bronze and maroon cardinalfishes (Apoffon nematoptei*us) , ready to 
retire into their stony sanctuaries at the approach of our staring 
camera lenses (pi. 11, fig- 1) . 

In the undercut shoreline of Koror nearby we found a small cave 
of the sort that permeates many of the limestone islands (pi, 6, fig, 1) . 



496 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

It was too small to crawl into very far, but in its dark and narrow 
recesses we found some alcyonarian corals (TeZesio) hanging from 
the roof, completely exposed by the ebbing tide. Elsewhere there are 
larges caves, some at water level and large enough to admit a boat, 
others equally large but completely submerged • In most of the major 
limestone islands there are large caves that have collapsed, forming 
lakes connected with the sea by subterranean passages. These salt- 
water lakes contain large gobies, mussels, and sedentary jellyfish 
(Cassiopeia)* 

One of our reconnaissance techniques was to cruise slowly along 
the coral slope in the Lenore, watching for changes in the appearance 
of the reefs to indicate interesting spots to be examined more closely. 
We followed the bay shore of Koror south from the little cave, around 
the so-called Arappu Peninsula (Ngalap), where the submarine cliff 
was almost devoid of corals and the only conspicuous organism was a 
large, sprawling, branched, pale pink sponge that looked white in 
20 or 30 feet of blue water. Even the stiff snaky a wire corals" 
(Antipatharia) that usually thrive on the cliffs were missing. We 
could find no explanation for the absence of corals there, for the 
current flow is better than it is in many other places, the water clearer. 

After we passed through the narrow strait called Kaki-suido 
(Oyster Pass) in the Japanese reports (Palauan name Ngerilduul), 
the situation changed, and we came upon one of the most interesting 
areas in Iwayama Bay, one that we visited and revisited, each time 
to find something new. Here the undercut was very deep and the 
foliage of the jungle-covered slopes above hung far down over the 
water, blocking the midday sunlight and producing an almost con- 
stant twilight- For only a few moments in the afternoon could a 
few rays of sunshine slip through before the shadow of island 29 
across the pass crept up to throw the waters into increasing darkness. 
A yawning cavern gaped in the cliff wall, entirely under water, its 
roof festooned with huge, netlike antipatharians ("black corals") 
hanging down like drapes. They were so flexible and so large that 
they could grow only in hanging position (pL 18, fig. 2). When 
fresh, their polyps were brilliant orange in daylight, but they ap- 
peared almost white in the murky blue water of the cave, 

ANIMAL PARTNERSHIPS 

Only on this half -lit slope did we find the whip -corals, Junceella; 
they grew here like tall, waving grass reaching out toward the light, 
with their tips drooping like buggy whips- They form the hub of an 
interesting association that we will describe in detail in a future paper* 
The most unusual member of this association is a little transparent 
pink goby, a new species allied to the genus Gottogobim* It is a tiny 



PALATJ BAYER AND HARRY-ROFEN 497 

fish, not much more than an inch in length, that clings to the stalks 
of the whip -corals. They may be pursued up one side of the coral 
and down the other, but rarely will they move to another coral even 
though it is very near. Our ingenious Rikrik found that by sliding a 
coral stalk through his fist, he could catch its fish in a small net as it 
popped off the end. The water was often murky and the light always 
poor, but we did get a picture of Rikrik catching gobies in a field of 
junceellas (pi. 16, fig. 1) . Also clinging to the whip-corals were many 
handsome gray feather-stars, or stalkless crinoids (comatulids) , some 
of them grasping several corals at once (ph 16, fig. 2). The cirri (the 
clawlike "feet") of the crinoids irritated the surface of the corals and 
scarred it permanently wherever they had clung, indicating that these 
"free-livinsr" crinoids move about little, if at all. One of the crinoid's 
distant cousins, a bright orange brittle-star, lived entwined in the arms 
of the crinoidj and among them could also be found a spider crab, 
Harrovia, that had joined the partnership. Another crustacean, a 
small porcellanid crab, scuttled over the surface of the whip-corals, 
completing this curious association of vertebrates and invertebrates. 

Among the most studied of symbiotic relationships to be found on 
tropical reefs is that involving several species of large sea-anemones 
that allow certain kinds of small but colorful damselfishes (genera 
A?7ipkiprion and Dasoyllus) to seek protection among their tentacles, 
These clownfishes ? as they are sometimes called , rarely stray far from 
their host anemone, and are ready to dart down among the stinging 
tentacles at the first hint of danger (pL 14, fig. 2). In spite of years 
of study, the details of this association are still not clear (Gohar, 1948 ; 
Gudger, 1946). It is believed that the fishes avoid being stung by 
swimming in a distinctive fashion that is "recognized" by the coelen- 
terate- Clownfishes have been seen to drag food to the waiting ten- 
tacles of the anemones, but, on the other hand, we watched an Amphi- 
prion seize a tentacle of its host in its mouth and with a few quick 
tugs pull it loose and eat it. This finny ingrate expected, and received, 
sanctuary from the very anemone it had been nibbling upon, for it 
dashed headlong among the tentacles when we approached too closely. 
Some investigators have suggested that by eating bits of the anemone, 
AmpkipHon builds up an immunity to nematocyst poison, but this 
suggestion has never been scientifically confirmed. It does seem fairly 
certain, however, that the clownfishes recognize their preferred species 
of host anemone partly by sight and partly by chemical emanations. 
There is also some indication that the anemones do not sting their 
partner fishes because of some kind of chemical "recognition." 

Coelenterates and echinoderms seem often to play the host role in 
these partnerships, probably not because of any inherent good nature 
or native generosity, but because they are slow moving or sedentary 



498 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1056 

and therefore an easy mark. One of our most startling discoveries 
of the summer was a fish that lives with an echinoderm. This, of itself, 
is not unusual, for the eel -like pearlfishes (Campus^ Fierasfer^ and 
Jordanians) have long been known to inhabit the cloaca of holo- 
thurians (sea-cucumbers) and the body cavity of Gulcita (the cushion 
starfish). But our fish was a previously unknown species of clingfish 
(Gobiesocidae) and the echinoderm host was a feather-star (Coman- 
thw)) a completely unexpected combination- The clingfish was black 
with a bright yellow stripe down each side, perfectly camouflaged 
among the arms of its host. 

It was on August 7 that we discovered it, as we were returning from 
a 2- day trip to Ngemelis* We had stopped in Meharehar, the labyrin- 
thine lagoon of Eil Malk, to look for future collecting sites and to ob- 
tain some samples of the lagoon bottom sediments. It was a stormy 
day with heavy downpours that had hampered our observations and 
dampened the spirits of everyone aboard the Lenore* We had taken 
the bottom samples in the rain, and were heading for home by way of 
the inside route west of Urukthapel when we found ourselves over some 
coral flats near Ankosu Point, the southernmost cape of Urukthapel. 
Not wishing to pass up any likely localities, we dropped anchor and 
went over the side to look around. The water was about 6 feet deep 
and the bottom was covered with a tangle of staghorn coral 
(Acropora)) most of it lying loose upon the sandy bottom* A few 
knolls of massive coral could be seen, with chalice-shaped acroporas 
and sea-fans (Afelithaea) growing on them. Here and there, hidden 
among the corals, we found a many -rayed spiny starfish (Acanthaster 
planci) , which is a real danger to bare feet (even those tough enough 
to disregard the jagged coral). It is widely feared by the natives of 
Micronesia, and with ample justification. A friend of ours was vir- 
tually incapacitated for a week or more by wounds inflicted by this 
animal, and he was not fully recovered for a month or more* 

Between the coral branches everywhere, and on the coral knolls, 
the restless, fernlike arms of feather-stars swayed with the rising 
tide. Because we knew that crinoids are ever-gracious hosts to a va- 
riety of invertebrates, we collected some of them to find their lodgers. 
There are usually two kinds of shrimp, a galatheid (or "squat-lob- 
ster"), and a polychaete worm, all protectively colored to match their 
host — usually black and greenish yellow in this locality. The first 
feather-star that Adair Fehlmarni collected had some blaek-and- 
yellow striped shrimps among its arms. Safely inside a glass vial, 
they gave us a real surprise* They were not shrimps at all, but 
fishes — and clingfishes at that — the only ones we would find all sum- 
mer. A careful search disclosed a number of additional specimens 
before the current became so swift that we could work no longer. We 



PALAIT — BAYER AND HARRT-EOFEN 499 

continued our homeward trip with one of the prize catches of the 
expedition — some tiny black-and-yellow fishes little more than an inch 

in length. 

In practically any protected sandy area of the lagoon we could ex- 
pect to find the unusual partnership of shrimps and fishes that we 
first observed at If aluk Atoll in 1953. We were delighted to find this 
association at Palau in such shallow water and in such abundance 
that we could observe it closely and collect the animals in numbers* 
At Palau, two kinds of alpheid shrimps and at least four kinds of 
gobies live together with identical habits. 

Each pair of shrimps excavates its own burrow and then plays 
host to a pair of fishes. The fishes sit just outside the mouth of the 
shrimp burrow (pi 14, fig, 1) while the shrimps repair and deepen 
it, bringing load after load of sand to the surface on their large claws, 
bulldozer-fashion. But should any danger threaten, the gobies dart 
down the hole in a trice, tumbling the shrimps over in their haste. 
From this activity the shrimps detect that something is amiss and 
cease digging until the gobies regain their composure and their usual 
position on the front doorstep. Whether this inadvertent warning 
is the only benefit derived by the shrimps from the association we are 
unable to say at present- It seems probable that the gobies obtain, 
in addition to shelter, an occasional banquet at the expense of their 
hosts, since one of the specimens collected was stuffed with larval 
crustaceans — probably the young of the shrimps whose home it had 
shared. 

There is still much to be learned about these and various other rarely 
observed biological associations, and it will take patient observation, 
study, and experimentation in the field before we know the exact 
nature of the relationship between the partners and how it may have 
developed. 

THE OUTER REEFS 

The outer reefs have an entirely different appearance from the 
lagoon reefs of Iwayama Bay and the staghorn coral thickets of the 
shallow passes- On the west side of the archipelago the reefs are 
barriers, but on the east they are fringing reefs that follow the land 
closely j with an offshore barrier in only a few places, Whether barrier 
or fringing reefs, they are bathed with the always clean water of the 
open sea and pounded by its sometimes thunderous surf. Different 
and stronger corals live in these exposed situations, and a whole new 
population of fishes swims among their branches. The surgeonfishes, 
butterfiyfishes, and wrasses that live here are for the most part 
peculiar to this zone of churning, turbulent water. Few species of 
the quiet lagoon waters are hardy enough to adapt to this rigorous 
environment. Sharks and barracudas in particular prefer this region. 



500 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

constantly patrolling it for food. Certain surgeonfishes also are char- 
acteristic of the surf zone, among them Accmthurus UneahtSy a yellow 
and neon-blue striped fish that loves turbulent, surging water* Their 
relatives, the fan-finned zebrasomas, hide under the spreading table 
corals. Huge parrot fishes, six or more feet in length, graze like 
cattle upon the corals, producing a noisy chorus as they crunch their 
limy meals in large, gregarious schools, Moray eels [Gyrrmw thorax) 
lie hi wait in their holes, their malicious eyes alert for any unwary 
fish that may pass within striking distance. The butterflyfishes 
(Chaetodon), so named because of their brilliant coloration, are 
especially numerous and are endowed with an insatiable curiosity. 
They seem unafraid of divers and will approach very close in order 
to get a good look. On many occasions they have startled us by a 
quick nip or a tug on some especially attractive hair. This has an 
electrifying effect upon swimmers already fearful of a surprise attack 
by prowling sharks, and the sheepish feeling combined with relief 
upon seeing these small challengers, is more amusing now than it 

was on the reel 

The spiny lobsters (Pamilirus) that lurk in crevices of the outer reef 
are different species from those found in the lagoon. So are the coral 
crabs (Trapezia) among the massive branches of pocilloporas, veri- 
table giants by comparison with their relatives in more protected 

waters. 

Any specimens taken from the surf zone are collected only through 
great exertion, for it is one of the most difficult habitats to sample- 
Swift currents rapidly dissipate the rotenone and sweep away the speci- 
mens killed by it. The collector himself may be picked up by the 
swell and dropped 15 or 20 feet away, as likely as not on a jagged 
cor ah When explosives are used instead of rotenone, all the nearby 
sharks materialize as if from nowhere, considering its sound a dinner 
bell inviting them to a free meah Needless to say, we used this explo- 
sive technique infrequently, so as to avoid becoming the piece de 
resistance of some shark's luncheon. 

Near the reef passes and in the deeper waters offshore the corals 
flourish in stony gardens of eerie and awesome beauty. Some species 
of Acropora produce towering spires and antlers (pL 7, fig. 1) that 
bring to mind the reefs made by the same genus of corals in the West 
Indies, and others form great, spreading disks and platforms com- 
posed of tiny branchlets, a growth form peculiar to the Indo-Pacific 
area (pi. 13). The sea gardens of swaying alcyonarians (sea-fans 
and sea-whips) that give the West Indian reefs their color and fluid 
beauty are nowhere to be seen, for the alcyonarians here are nearly all 
massive, rubbery kinds (pi. 13, fig. 2) that have not the elegance of 
their Caribbean relatives. The few species of sea- fans that do occur 



PALATJ — BAYER AND HARRY-ROFEN 501 

are more abundant along deep channels or in the protected lagoons 
than on the seaward reefs, 

EEL-GRASS ENVIRONMENT 

A conspicuous feature of the Palau lagoon is the great extent of 
sandy bottom covered with eel-grass (Enhalus acoroides and some re- 
lated species) . It is a distinctive and complex habitat. The most ex- 
pansive eel-grass beds lie near Peleliu, to the south of Koror, al- 
though the west coast of Babelthuap also has some fine ones. We 
made two trips to Peleliu to survey the grass beds there, but on both 
occasions we found poor conditions due to the stage of the tide. The 
water was so murky and full of plant debris that collecting was un- 
pleasant and photography impossible. 

Among the animal inhabitants, fishes are particularly abundant and 
thrive in the eel-grass environment Sharks, jacks, barracudas, and 
other predaceous fishes constantly search the eel-grass beds for prey. 
Food is not easy to find here, even in the midst of plenty, for the eel- 
grass forms the home of many highly specialized fishes that blend with 
their background in both form and color. Some of them, such as the 
pipefishes (Syngnathidae) and certain wrasses (Gheilio), are elongate 
in outline and green in color, so perfectly camouflaged that sharp eyes 
are needed to separate them from the grass in which they live. Others, 
such as the parrot fish (Scarichthys) , spinefish (Siganus), and some 
snappers (Lethrinus) 7 are not shaped like the grass blades but are so 
much like it in color that they are virtually invisible. The filefishes 
(M onae ant hus) ^hlermies {Petrosoirtes)^ and dragonets {OalMonymus) 
go a step f arther in having their bodies covered with waving filaments 
and hairlike growths that resemble the hydroids and other epiphytes 
covering the eel- grass blades. 

The Palauan boys pointed out some peculiar little black- an d-yellow 
fishes that were swimming about a water-logged branch half buried 
in the sand and hidden by grass and warned us that they w r ere very 
dangerous. When we insisted upon catching them, Sumang Y. and 
Rikrik must have regretted pointing them out to us. The fish were 
small catfish (Plotoms anguiUaris) with barbed pectoral and dorsal 
spines that are venomous and can inflict a nasty wound. They were 
swimming in a curious manner, very close together and wiggling vigor- 
ously, in a compact school that moved slowly forward like a dark 
cloud. It was simple to frighten them into the range of a large dip- 
net, and each fish captured caused our sturdy Palauans to wince in 
anticipation of the painful punctures to come when we pulled them 
out of the nets, and later, when we placed them in containers of for- 
malin, A month or so later, when we were in Japan to consult with 
biologists who had worked in Palau before the war, we encountered this 



502 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1056 

same fish in southern Honshu, It was living in a completely different 
ecological situation, in rocky tide pools near the Seto Marine Biologi- 
cal Station at Shirahama. 

One of the major eel-grass inhabitants is a mammal, the little-known 
dugong, or sea-cow {Dugong dugon)* During our stay in Palau we 
were always especially watchful while passing over eel-grass areas, 
in the hope of seeing some dugongs, but as they are quite scarce we 
never did see one in its natural habitat. During our last few days at 
Koror, how T ever, some Palauans speared and captured alive a half- 
grown specimen that provided us with considerable excitement and be- 
came the first ever to be exhibited alive in an American aquarium. 
We bought this dugong, otherwise destined for sale as food in the local 
market, and kept it alive in a large pool at the end of Koror dock 
until we were ready to begin our return trip to the United States* 
Then we caught it, wrapped it in wet blankets (ph 20, fig. 1), and 
carried it, lying on the floor of the plane between our seats, to Guam 
(pL 20, fig- 2). At Guam it was ensconced in a sturdy crate and 
transferred to a commercial airline for shipment to California, The 
California Academy of Sciences had arranged for its transportation 
to the Steinhart Aquarium, where it proved to be an unusually popular 
exhibit. It was certainly the first Palauan dugong ever to fly to 
America, and we have no doubt provided some material for the "talk- 
ing picture" carvings that decorate Palauan abais, 

HERPETOLOGICAL STUDIES 

Another Palauan animal that never before had been displayed alive 
in an American 200 or aquarium, and which we succeeded in bring- 
ing back to the United States, is the deadly poisonous sea snake 
{Laticauda colubHna)^ a relative of the cobras and coral snakes. 

Our first encounter with the banded sea snake in the wild was on 
a field trip to Ngemelis, a group of islands along the southwest barrier 
reef. We were making our way toward the beach just before dusk 
when, a hundred yards or so from shore, we came upon a huge snake, 
a good 6 feet in length, slowly working its way seaward along the 
bottom, poking its head into nooks and crannies and in and out of 
corals, carefully feeling with its tongue. It paid no attention to us, 
and we stood or swam near it for several minutes, in water perhaps 
5 feet deep- At no time did we see it surface for air, and as we con- 
tinued on our way it was still swimming seaward along the bottom. 

The Palauans have an odd story about using sea snakes to catch fish. 
They say that if one holds the snake by the tail, it will probe among 
the rocks and catch fish, which can then be easily seized- We never 
saw this method of fishing practiced- Probably no modern Palauans 
are courageous enough to try it, for our helpers invariably let out 



SMITHSONIAN REPORT, 1956. — BAYER AND HARRY-ROFEN 



PLATE 11 




1 



Graceful, spotted cardinalfishes (A pagan nematopferits) hover over the branches of a 
Parties colony that is their refuge in times of danger. This little fish, with its red spots 
on a bronzy ground color, and a dark-brown cummerbund, is one of the loveliest denizens 
of protected reefs. 




2. Huge, fragile chalices of Montipora foliosa dominate the reef slope in the innermost part 
of K Division of Iwayama Bay, They form an uncertain retreat for fishes like the small 
damselfish {Pomacenttus) swimming above them, 



SMITHSONIAN REPORT. 1956.— BAYER AND HARRY-ROFEN 



PLATE 12 




L The beautiful falls of Matal Eigad on the Ateshi River, in Ngardmau Municipality of 
Babel thuap. Note figure at foot of falls. The only fishes collected above the falls were 
eels, which can travel overland around obstacles insurmountable to other fishes, (Photo- 
graphed by H. A. Fehlmann.) 




2, H. Adair Fehlmann and Rikrik collecting in a small stream on Arekalong Peninsula of 
northern Babelthuap, Such a locality yields many gobies, and prawns of the genus 
Macrobrackium* 



SMITHSONIAN REPORT, 1956.— BAYER AND HARRY-ROFEN 



PLATE 13 




1. Great, circular plates of acroporas grow one above the other like modernistic buildings 

in this scene along a reef channel south of Ngarcmediu. 




2, The convoluted mass in the foreground is a massive, fleshy alcyonarian, Sarcophyton f as 
large as a bushel basket. This type of alcyonarian is typical of Pacific reefs, whereas in 
the Atlantic the bushy gorgonians dominate. 



SMITHSONIAN REPORT, 1956, — BAYER AND HARRY-ROFEN 




1. At left, a pair of speckled gobies sit outside the burrow made by the dark-banded snapping 
shrimps. At right, the banded gobies are living with a pale, fine-striped shrimp that is 
almost invisible against the sand. 




2. At left, a clownfish {Amphiprion) pauses above its host anemone. At right, a clownfish 
can be seen hiding among the tentacles of the anemone, which do not sting it. 






SMITHSONIAN REPORT. 1956.— BAYER AND HARRY-ROFEN 




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PALAU — BAYER AND HARRT-KOFEN 



503 



great yelps of fear whenever we caught sea snakes (or any other kind, 
for that matter), and their repugnance extends also to eels, perhaps 
with more justification. Another droll story, frequently seen on the 
carved w talking picture" boards, tells how the sea snake taught 
Palauans to fish with nets. In this tale, a Palauan woman, whose 
children were a banana tree, a pussycat, and a sea snake, grew weary 
of her offspring and told them to make their own way in the world. 
The banana tree pointed out the impracticality of this suggestion, at 
least from its rooted standpoint, and thus won a reprieve, but the two 
more active "children" were sent packing. As they swam toward 
Orukuizu, the cat riding on the snake's head, they grew hungry, and in 
Palauan carvings we see the snake encircling some fishes so the cat 
could catch them as they swam out through the narrow space between 
the snake's head and tail. A villager who was up in a palm tree 
gathering toddy caught sight of this procedure and at once recognized 
its possibilities. He rushed down to the shore and asked the snake to 
show him how to fish that way, so the snake showed him how to set 
up a fence of sticks in shallow water, in which fish would be trapped. 
But the Orukuizu man still needed the cat to catch the fish as they 
passed through the mouth of the weir, so he asked the snake if he could 
have the cat, too. The snake agreed but made the man promise to give 
the cat one of the fish each time they made a successful catch. Such 
was the origin of cats and fishweirs* 

This particular species of sea snake is not so completely aquatic as 
are some of its relatives, and it could be found on the seaward shore of 
exposed islands, coiled up in underbrush on dry land or even in the 
branches of trees (ph 19, fig. 2) ♦ They were very common at Nardueis, 
off the southeast shore of Babelthuap, where we caught as many as a 
dozen in a couple of hours. Some of these are still living in the Stein- 
hart Aquarium in San Francisco, but the specimens destined for the 
National Zoological Park in Washington perished during the flight 
across the continent. 

The herpetological collections we assembled contain at least four 
other species of snakes, including a burrowing form {Typhlops) no 




The story of the sea snake, the cat, and the banana tree. Adapted from an actual Palaun 

carving, 



412570 — 57 SS 



504 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

larger than a moderate-sized earthworm- Geckos (nocturnal lizards 
with adhesive toe pads that aid in climbing) of several kinds abound 
in the trees and on window screens of buildings where they catch in- 
sects attracted to electric lights at night. The toad is extremely 
abundant in the vicinity of Koror, and two kinds of frog seem to 
be widespread throughout the islands. The monitor lizard is not 
especially common but may be encountered on Babelthuap. The 
collections and notes of Dr. Masamitsu Oshima, who studied the rep- 
tiles of Palau, were unfortunately destroyed during the course of the 
war before they could be published. Our collections will therefore 
fill an important gap in our knowledge of the terrestrial fauna of the 
Palaus. 

HOMEWARD BOUND 

By the time November rolled around and Project Coral Fish had 
been active in Palau for four months, we began to make plans for 
the homeward journey. On November 4 we made our last marine 
collecting station, and on the following day the dismantling of the 
laboratory began. All equipment had to be either packed for ship- 
ment or stored for use by Project Coral Fish II. Outboard motors 
were cleaned and boxed, the Lenore was stripped and crated ; collec- 
tions were packed in tins, drums, and boxes, labeled, and stacked up 
to await the next freighter to Guam. In a few days the laboratory 
was starkly in order and deserted. Arrangements were made for 
the transportation of the live cargo of sea snakes and dugong. "Plane 
Day" on November 15 was a typical rainy -season day in Koror. Rikrik 
and Sumang Y. went early to the dock and caught the dugong, and 
it was difficult to say if it was wetter in the pool or on the dock. 
Dugong, snakes, other precious collections, and ourselves were loaded 
on the Albatross amphibian in the still-pouring rain, and we were 
off, it seemed after having scarcely arrived, so swiftly did the months 
pass on these coral-fringed islands and broad lagoons. Only yester- 
day, so it seemed, we had made our way up one of Babel thuap's 
rivers as the crocodiles slid off the banks into the dark water, 
or had been listening to the little bird with a "pipe sweet as a 
flageolet" as we collected in some forest-bound coral bay. Perhaps 
nowhere else in the world are reefs and jungles so intimately asso- 
ciated that one can stand among living corals and at the same time 
collect many different kinds of orchids from overhanging branches. 
It will be many a year before the complexities of this multifarious 
habitat are thoroughly understood, but we think that Project Coral 
Fish is making a long stride toward that understanding and, there- 
fore, toward a better understanding of life in the seas as a whole. 

During the 1955 expedition to Palau, Project Coral Fish assembled 
thousands of specimens from 276 collecting stations extending from 



PALAU — BAYER AND HARRT-EOFEN 505 

the northern end of Babelthuap to Peleliu. Phylogenetically, the 
collections contain animals ranging from the Protozoa to the verte- 
brates, as well as many specimens of marine plants. The largest 
collections are those of fishes, crustaceans, and sponges, but the smaller 
collections are no less significant. Taken together, they represent a 
contribution toward a more complete faunistic and zoogeographic 
knowledge of the Palaus, an especially significant one in view of the 
fact that the collections made in the same territory by Japanese 
scholars prior to World War II are either widely scattered over Japan 
or were destroyed during the war. In addition to their systematic and 
zoogeographic value, these collections include information on reef 
communities and biological associations, which forms the framework 
of reef ecology. In spite of its encouraging success, the first field 
season of Project Coral Fish did not accomplish all there was to be 
done in Palau. Neither will the 1956 season. Some of the smaller 
component projects will certainly see early completion, but they bring 
into bold relief many other problems, not a few of which demand 
an experimental approach that can succeed only at a place like Palau. 

CHARACTERISTICS OF THE PALAUAN MARINE FAUNA 



The large collection of marine animals assembled in Palau during 
the summer and fall of 1955 is only now being studied in detail, 
so it is not yet possible to analyze the fauna from a zoogeographic 
standpoint However, some impressions of both its richness and its 
affinities were inescapable during the course of observing, collecting, 
and preserving the many specimens handled. These impressions, 
superimposed upon what we already know from the literature, enable 
us to reaffirm the East Indian relationships of the Palauan marine 
fauna. 

Many East Indian fish groups not common in the oceanic islands 
were found to be abundant in the Palaus. Especially noticeable is 
the archerfish (Toxotes)j which travels in large schools along the 
lagoon shores, particularly in mangrove regions, where it penetrates 
the narrow waterways and ascends a considerable distance up streams 
and rivers. Glassfishes (Ambassis) and spotted scats {Scatophagies) 
are abundant in the mangrove swamps and lower reaches of streams 
but are not found in the limestone islands of the southern Palaus. 
Fresh- water eels {Anguilla) grow up to 5 feet long in even the 
smallest streams. Some groups of fishes, such as the snappers (Plecto- 
rkynchus, CaestOj Lethriniis, and others) are represented by many more 
species than are known from the multitudes of oceanic islands in the 
Marshalls, Marianas, and Carolines, 

Among the invertebrates, the only two shallow- water gorgonaceans 
(Octocorallia) to extend an appreciable distance eastward into Micro- 



506 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1656 

nesia, Suftergorgia mollis and Rumphella aniipathes, are common 
East Indian species that occur also, as expected, in the Palaus. An- 
other species of Subergorgia, two magnificent nielithaeas, the so-called 
"Plexaura" ffava^ a species of Euplexaura^ and Junceella fragilis are 
more or less common in Palau, indicative of close ties with the East 
Indies and Philippine faunas. Among the Alcyonacea, De?idro- 
nepkthya occurs in abundance and Studeriotes was found in slightly 
deeper water in the lagoon, both genera being well represented in the 
Philippines, These various octocorallian genera are centered mainly 
in the East Indies-Philippines area, but range also northward to 
Japan and southward to Australia, thence eastward to Fiji, Tahiti, 
and Tonga, but reduced in species peripherally* Furthermore, the 
alcyonarian genera typical of Micronesian reefs, such as Sinularia^ 
Sarcophytorij Lobophytmn^ and SpkaereUa y are represented in Palau 
by a greater number of species than is usual in the central Pacific 
atolls* 

The molluscan fauna is notably richer in many elements than that 
of the Micronesian atolls to the east, especially in such genera as 
Mur&x, Spondylus, and Peeten. Since the distribution of Shnnia and 
related gastropods is tied to that of the gorgonians on which they live, 
they accordingly do not extend very far to the east, probably not be- 
yond Palau. However, the large egg-cowry (Ovula ovtim) y 
which lives on soft-corals, enjoys the much wider range of its hosts. 

The fauna of the Palaus will probably not be found to approach 
the richness of that of the Philippines, East Indies, and New Guinea, 
but it certainly far overshadows that of the oceanic atolls. 

PROSPECT 

As we write these words, Project Coral Fish II is continuing the 
program of investigations embarked upon in 1955. It is anticipated 
that the field phase of a survey of the fresh-water fish fauna of Babel- 
thuap, begun in 1955, will be completed successfully during the pres- 
ent season, and the marine survey continues with special attention to 
areas not previously visited. The research program has been broad- 
ened by the addition of an oceanographer to the field party, so that 
more detailed data regarding the aquatic environment may be 
gathered. 

During Coral Fish III in 1957, we hope to complete our ecological 
resurvey of Iwayama Bay, and to make more detailed studies of some 
of the commensal, symbiotic, and parasitic relationships mentioned in 
the foregoing pages. The results of the 1956 expedition should permit 
much more efficient observations on the physical environment and en- 
hance the value of the Iwayama resurvey. We hope that it will also 



PALAU — BAYER AND HAKRY-ROFEN 507 

be possible to inaugurate a program of current and plankton studies, 
especially within Iwayama Bay, to determine the reasons for, and to 
provide a better understanding of, the distribution of various corals 
and other benthic invertebrates within the confines of the Bay. 

ACKNOWLEDGMENTS 

The fieldwork described in these pages was carried out under ar- 
rangements with the Pacific Science Board (NAS-NRC) and the 
Office of Naval Research [contract N7onr-291(57) J, Work in the 
Palaus would have been impossible without the approval and thorough 
cooperation of the Trust Territory of the Pacific Islands, Especially 
we wish to express our appreciation to High Commissioner D. 
H, Nucker, Executive Officer A. M« Hurt, and their staffs in Guam. 
In Palau, District Administrator Donald Heron extended every pos- 
sible courtesy, and his staff, including Francis B* Mahoney, Robert P. 
Owen, and many others, were wholeheartedly cooperative and inter- 
ested in our program, We likewise cannot overlook the Trustees of the 
George Vanderbilt Foundation and the authorities of the Smithsonian 
Institution, who made our participation j>ossible. 

PALAUAN GEOGRAPHIC NAMES 

The names used in this account are those to be found on the charts 
of the U. S* Hydrographic Office and the maps of the U. S. Corps of 
Engineers. The spellings of these names may bear little resemblance 
to their pronunciations. We therefore offer the following table, the 
left-hand column listing the names as taken from published charts and 
used herein, and the right-hand column listing the phonetic spellings 
by Kramer, or supplied to us by Suxnang Y. 

Angaur ( Island )„„ » ._. _„___ a Ngeaur 

Aiikosu (Point)- — .„« Nghus 

Arakabesan (Island), . . „_ Ngarekobasfing 

Arekalong ( Pen insula )__— ._. .„_ Ngaregolong 

Ateshi (River) [also Addeido] a Did 

Aulong (Island) ._„_ a Ulong 

Aurapusbekaru (Island) [also Auluptagel]- Ulupsagel 

Babel thuap (Island) _ „_ Babldaop 

BilMalk (Island) __ a Ilm&lk 

Geruherugairu (Pass) „_ „_ Ngaregelng&e! 

Kayangel (Atoll) — . , „„__„„_ Nggeiangel 

Malakal (Island) -„. Maldgal 

Matal Eigad (Palls of a Did R.) Madal a Jegad 

Nardueis (Island) . Ngarduals 

Orukuizu (Islands) _«„ , ____ Ngerkuid 

Palau [also Paiao, Pelew, etc.]™* Pelan or Belau 

PeleUu ,„__„_ Pelfiiou 

Urukthapel — — __, ,„ „, ., _. Ngurukd&pel 



508 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

LITERATURE 
Abe, Noboru. 

1937. Ecological survey of Iwayama Bay, Palao. Palao Trop. BioL Stat 
Stud., voL 1, No. 2, pp. 217-324, 42 figs. 
Abe, Noboru ; Eguchi, M. ; and Hmo, F. 

1937, Preliminary survey of the coral reef of Iwayama Bay, Palao, Palao 

Trop, BioL Stat Stud., vol. 1, No, l f pp. 17-35, 1 text fig., 2 pis., chart 

Abe, Tokihabtl 

1939. A list of the fishes of the Palao Islands, Palao Trop. BioL Stat Stud., 
vol. 1, No. 4, pp. 523-583, 
Davenport, Demobest, 

1955. Specificity and behavior in symbioses. Quart. Rev. BioL, voL 30, 

No, 1, pp. 29-46, 7 figs. 
Gohar, EL A* F. 

1948. Commensalism between fish and anemone (with a description of the 
eggs of Amphiprion Mcinctus Riippell), PubL Marine Biol. Stat 
Ghardaqa (Red Sea), Egypt, vol. 6, pp. 35-44, 4 pis. 

GUDGEE, E. W. 

1946. Poinacentrid fishes symbiotic with giant sea anemones in Indo-Paclfic 
waters, Journ. Roy. Asiatic Soc. Bengal, Science, voL 12, No. 2, 
pp. 53-76, 2 pis, 
Habby, Robebt R. 

1956. "Eugenie" the dugong mermaid. Pacific Discovery, vol. 9, No, 1, pp. 

21-27, 5 figs. 
Keate, Geoege* 

1788. An account of the Pelew Islands, situated in the western part of the 

Pacific Ocean. Composed from the journals and communications 

of Captain Henry Wilson, and some of his officers, who, in August 

1783, were there shipwrecked, in the Antelope, a packet belonging to 

the Honourable East India Company. London. 
Kramer, A. 

1917-1929, Palau, Ergebn. Siidsee Exped. (Thilenius, ed.). II. Ethno- 

graphie, B, Mikronesien, vol. 3. PL 1, 1917 ; Pt 2, 1919 ; Pt. 3, 1926 ; 

Pt 4-5, 1929, 

KlJBAEY, J. S. 

1873, Die Palau Inseln in der Siidsee. Journ. Mus. Godeffroy, vol. 1, No. 4, 

pp. 177-238. 
1895, Ethnographisehe Beltrage zur Kenntnis des Karolinen Archipels, 
Leiden, 
Nanyo Gunto Buwea Kyokai (South Seas Cultural Association)* 

1938. Nanyo Gunto Shashin Cho (Photo album of the South Sea Islands), 

Tokyo. 



The following text is generated from uncorrected OCR. 
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Project Coral Fish Looks at Palau 

By Frederick M. Bayer 

United States National Museum 
Smithsonian Institution 

and 

Robert R. Harry-Rofen 

The George Vanderbilt Foundation 

[With 20 plates] 

We who live on continents can rarely appreciate the vastness of 
the world's oceans. Those of us who may be prompted by business or 
pleasure to traverse them learn of their two-dimensional magnitude, 
but there are a few of us who are privileged to investigate the secrets 
of the seas first-hand, by living among them. We look upon the seas 
in their role as an environment and seek to unravel the interwoven 
facts of life within them. One of the first facts we learn is the com- 
plexity of their many-faceted wonders, and we consider ourselves 
fortunate when we are able, as it were, to polish a few of these facets 
so as to see more clearly into them. 



Of all the seas, the one with the greatest area, greatest depth, and 
most to tell us is that restless giant, the Pacific. Even before we begin 
to study it we must concede, if not defeat, at best a draw, for it is prac- 
tically axiomatic that one solution leads to another problem. When 
our words are as antiquated as those of the Renaissance pioneers Belon 
and Rondelet and Marsigli now seem to us, people will still be learning 
new things about this watery one-third of our planet. 

When we look upon the Pacific, or any ocean for that matter, as an 
environment, the central problem deals with the physical and chemical 
properties of the fluid medium that make it adequate to support life. 
This is a vast field of investigation to which many people in many 
laboratories are devoting tireless efforts. We, however, as biologists, 
can devote time to such problems only when they have direct and im- 
mediate bearing upon some question involving the organisms with 
which we are concerned, and even then we must rely upon specialists 
in those restricted fields for most of the information we need. The 



481 



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482 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

next problem is a qualitative one regarding the population of the sea 
what are these creatures that populate the waters? This is a basic 



problem in marine biology ; upon it depend the solutions to problems 
of economic, or ecologic, or purely biologic interest. We must know 
what organisms wo are working with before we can determine how 
they live together in communities, how they depend upon one another, 
and how they affect us. 

Creatures of the sea do, in fact, affect the affairs of man in many 
ways. Many of them have served us for food since the beginning of 
mankind. When we build structures in the sea for our own purposes, 
certain of the animals and plants whose domain we have invaded use 
those structures to their own ends and thus either destroy what man 
has made or so befoul and beclog it as to render it worthless. When 
we sail in tropic waters, other marine life — corals and algae — has been 
there long before us and raised up an edifice that passively awaits the 
unwary navigator and his fragile keel. If we are thrown into the sea, 
or when we voluntarily venture into it, still others may unintentionally 
do us bodily damage or even deliberately seek us out as a meal. Most 
"dangerous" of all, to marine biologists anyway, are those that have 
such bizarre or complicated ways of life that they entice us to devote 
most of our lives to learning of them and solving their riddles. 

Since the close of World War II, interest in the Pacific Ocean has 
been increasing steadily. A number of expeditions were sent out to 
study the tropical Pacific, among which were those of the Pacific 
Science Board (National Academy of Sciences-National Kesearch 
Council) and the George Vanderbilt Foundation. Among the 
expeditions of the Pacific Science Board were those comprising the 
5-year Coral Atoll Program, in some of which the present authors 



participated. 

The expeditions of the Coral Atoll Program did much to broaden 
our knowledge of life on the coral atolls of the Marshalls, Gilberts, 
Carolines, and Tuamotus. But field team studies came to a close 
before the most interesting part of the Pacific could be studied : the 
western rim, the f aunal gateway to that vast coral world that reaches 
to Hawaii, the Galapagos, and even our own western shores. Moving 
eastward from the Malay Archipelago and its wonderfully rich fauna, 
we find no depletion through the Philippines, but what of the western- 
most islands of Micronesia? They are a scant 600 miles east of the 
Philippine Islands (see map, fig. 1 ) , no journey at all for sea creatures 
with free-swimming young stages. However, between the Philippines 
and the Palau Archipelago lies one of the greatest deeps in all the 
seas. How many of the East Indian species have been able to span 
this deep? Even the submarine ridge upon which the Palaus are 
situated, extending northeastward from the Moluccas, is covered by 



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PALAU — BAYER AND HARRY-ROFEN 



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484 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

waters too deep to serve as a shallow-water passageway for littoral 
forms. Are the Palaus now so isolated that special endemic species, 
found nowhere else, have evolved from their East Indian ancestors? 
How do the reefs, the habitat of most tropical shallow-water animals, 
as we see them in Palau, differ from those elsewhere in Micronesia, 
and from those in the East Indies ? Such were the questions we hoped 
to answer when we, as part of a 4-man team, set out for the Palau 
Islands late in June 1955. 

PROJECT CORAL FISH 



Together, we formed Project Coral Fish, a continuing field program 
devoted to the study of the marine biology of the high islands and 
atolls in the Trust Territory of the Pacific Islands. This program was 
initiated in 1954 by the George Vanderbilt Foundation at Stanford 
University, with the cooperation and support of the Pacific Science 
Board (National Academy of Sciences-National Research Council), 
the Office of Naval Research, the United States Department of the 
Navy, the United States Trust Territory of the Pacific Islands, and 
the Smithsonian Institution. Aside from the present writers, the team 
included H. Adair Fehlmann, assistant curator of the George Vander- 
bilt Foundation collections at Stanford University, and Sterling H. 
Pierce, technical assistant. We hoped to gather data and specimens 
of all kinds to give us an approximate answer to the question : What 
lives in the waters of Palau ? With this basic information we would 
be in a better position to answer such practical questions as: What 
kind of marine life can be exploited for food ? How can their supply 
be conserved? What other marine products of economic importance 
could the Palauan people develop successfully ? What are the dangers 
in fishing the reefs, and how can they be avoided ? 

As a byproduct of our basic task of finding out what animals popu- 
late the reefs of Palau, we studied the communities living in various 
kinds of habitats, especially in Iwayama Bay, and the strange associa- 
tions that develop between different kinds of animals. 

After many months of preparation, during which supplies and 
equipment were assembled and shipped to the western Pacific, the four 
team members assembled on June 22, 1955, at the George Vanderbilt 
Foundation headquarters on the Stanford University campus, and 



final plans for the trip to Palau were made. Under orders from the 
Chief of Naval Operations, the following day we left Moffett Field, 
Calif., aboard a military transport plane bound for Guam. 

Two days and more than 6,000 miles later, we arrived in Guam, 
where we learned that the expedition equipment had all been for- 
warded to the Palaus on schedule. D. H. Nucker, the Acting High 
Commissioner of the U. S. Trust Territory of the Pacific Islands, with 



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PALAU — BAYER AND HARRY-ROFEN 485 

whom arrangements for the expedition had been made months pre- 
viously, issued the permits necessary for our travel and field work in 
the islands. All was in readiness. We were in the Tropics again, and 
our destination was almost in sight. 

It was an impatient group that awaited the next weekly flight of 
the Transocean Airliner to Koror, but at last Thursday came and 
we were at the air terminal in the Naval Air Station, Agana, early 
that morning. A few moments after Miss Thelma Gorman, of Trust 
Territory Headquarters, had bustled us and the other passengers 
aboard the Albatross amphibian and breathed a sigh of relief, we were 
airborne. 



The islands of Palau lie some 800 miles southwest of Guam, a flight 
of about six hours, including stops at Ulithi Atoll and Yap. At 
Ulithi, we touched down on the airstrip to discharge passengers, 
where two years previously we ourselves had landed on our way to 
Ifaluk, a tiny jewel in the sea that captured our hearts as could no 
other spot. But that is another story, and we were back in the air 
before we could reminisce about it. At Yap, the lagoon is the air- 
strip and we made a water landing to discharge passengers, cargo, 
and mail. Again, after a pause of only a few minutes, we were taking 
off on the last leg of our journey halfway around the world. The 
next land we would see would be Palau — first the great, hilly island 
Babelthuap, then Koror with its settlement and the broad harbor on 
which we would land. Reaching up toward us from the sea below, 
were the jagged ridges of the islands we would come to know so well 
(pi. 1 , fig. 1 ). Amid sheets of spray we settled in the green water of 
the lagoon and taxied up to the ramp that led the plane out of the 
water. At last we were in Palau. 

THE PALAUS 

Although the Palau Islands were discovered in 1710 by Francisco 
de Padilla and his pilot, Joseph Somera, on the galleon La Santisima 
Trinidad (and may have been sighted earlier, perhaps by Diego de 
Rocha in 1525-26 or Lope Martin in 1566), the first Europeans to 
publish an extensive account of their visit were the crew of the 
British East India Company's Antelope in command of Capt. Henry 
Wilson. The keel of the luckless Antelope struck the coral rocks 
of the barrier reefs near Aulong Island on the morning of August 1 0, 



1 783. Only one man was lost in the disaster, and the remainder of the 
crew escaped safely to set up camp on Aulong. The castaways salvaged 
every usable item from the wreck and, although the Antelope was a 
total loss, were able to build another vessel large and seaworthy enough 
to take them all safely to Macao. This vessel, the Oroolong, brought 
700 Spanish dollars (about equivalent to the U. S. dollar of the 



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486 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

-S A SS\JJ,. loud 

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PALAV ISLANDS 

Figure 2 — The Palau Islands. Adapted from U. S. H. O. chart 6073. 



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SMITHSONIAN REPORT, 1956 —BAYER AND HARRY-ROFEN 

Koror from the air, with Iwayama Bay and the limestone islands in the distance, 
abrupt headland on the horizon at right is Ngaremediu Point on Urukthapel. 



The 



Looking east through Ngasaksao Pass, the eastern entrance to Iwayama Bay. To the 
left is Ngalap region of Koror, and to the right Kwannon (Ugeliungs) Island, with conical 
Ngaraglbukl (Ngergelbakl) Rock in the pass. 



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SMITHSONIAN REPORT. 1956. — BAYER AND HARRY-ROFEN 

1 . The Lenore in Lebugol Passage, the narrow western entrance to Iwayama Bay. This 
boat was used by field parties to reach the more remote localities investigated in the 
Palaus. 

»„',![ Ti"'iifi|i|iiw 



. A building that was once part of the Japanese meterological station on Koror now accom- 
modates biological activities in the Palau District of the U. S. Trust Territory. The 
ground-floor laboratory to the left of the main entrance is occupied by the George Vander- 
bilt Foundation, and the apartment above (now remodeled) provides living quarters for 
members of its field parties. 



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SMITHSONIAN REPORT. 1956 — BAYER AND HARRY-ROFEN 



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1 . One of the authors preparing to descend in a self-contained underwater breathing 
apparatus (aqualung). 

2. A dark green feather-star (Comanthus) of the species that is the host of the commensal 
clingnsh. The small urnlike objects to the left of it are colonies of an ascidian, Didemnum 

Igfis 
ternatanum. 



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SMITHSONIAN REPORT. 1956.— BAYER AND HARRY-ROFEN 

1 . Shallow reefs in Iwayama Bay are in some places dominated by the large poritid coral 
Goniopora. The stubby branches of its skeleton are obscured by the exceptionally large 
polyps, which remain extended even in daylight. Unlike most anthozoan corals, it can 



sting, thus making collecting near it very unpleasant. 

This species of mushroom coral, Fungia actiniformis palawensis, has long, creamy-yellow 
tentacles that completely hide its stony skeleton which is nearly the size of a saucer. It 
is a solitary polyp that never forms a colonial skeleton as most other reef corals do. A 
small specimen of a wrasse, Cheilinus, that is common in enclosed bays like Iwayama, can 
be seen at the top of the picture. 



[Begin Page: Page 487] 

PALAU — BAYER AND HARRY-ROFEN 487 

period) when sold in Macao, and stands as a tribute to British forti- 
tude and resourcefulness in the face of adversity. 

The English, although at first apprehensive, found no difficulty 
in establishing friendly relations with the Palauans. Even though 
they "had not on board philosophers, botanists, draughtsmen, or gentle- 
men experienced in such scientific pursuits as might enable them to 
examine with judgment objects which presented themselves, or trace 
nature through all her labyrinths," they nevertheless gave us a fasci- 
nating account of the islands and people and their experiences among 
them. 

Palau and the Palauans today present a far different impression 
from that described by Wilson and his men in 1783. Although the na- 
tives had complete power over the English, they did not take advantage 



of their superior position, but instead did everything possible to help 
their hapless visitors. Admittedly, the Palauans were awed by fire- 
arms, and the English assisted Abba Thulle (Ebadul or Ibedul) the 
chief of Koror, in some of his interisland campaigns. Good relations 
were thereby strengthened, but it probably was not guns alone that 
made the Palauans friendly. "We ourselves have been residents of 
islands where the people have had little real contact with civilization 
as we know it, and found that they remain much as Wilson pictured 
the Palauans, with a high regard for honesty and respect for their fel- 
lows. Now the Palauan people have become sophisticated and mun- 
dane after 20 years of German administration and another 26 under 
the Japanese. Missionaries long ago insisted that they give up their 
native way of dress (or undress) in favor of less practical European- 
style clothing. Their airy, thatched houses and abais, or men's houses 
(pi. 3, fig. 1), decorated with colorful murals depicting historical and 
mythological events or droll folk tales, have given way to quonset huts 
and quasi- Japanese frame structures. Their elegant outrigger canoes 
are becoming a rarity, replaced by dirty little diesel boats like the 
My Flower — anything but flowerlike — that plies between Peleliu and 
Koror. 

The Palau Archipelago in the western Caroline Islands (fig. 2) ex- 
tends northeastward from about latitude 6°53' N. to 8°06 / N., or 
over 70 miles, at a longitude of about 134°29' E. The main islands, 
dominated by Babelthuap, lie between latitudes 7° and 7°45' N. Ex- 
cept for Kayangel, the northern part of the islands is volcanic in 
origin. Babelthuap is about 25 miles long and 8 miles wide, covered 
with rugged hills and dense jungle, rolling grasslands, sparkling 
streams and dashing water falls, (pi. 12, fig. 1) , and fringed with man- 



grove. Twenty miles to the north of it is Kayangel, a true atoll like 
the islands far to the east, and to the south of it is the maze of lime- 
stone ridges and conical islets (pi. 1) that forms one of the most re- 

412575—57 32 



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488 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

Sketch of old abais, based on several published photographs. 

markable features of Palau. Here and there on the edge of the reefs 
there are islets of wave-tossed coral debris and sand, much like the 
atoll islands. Together, the Palaus offer the most diversified collec- 
tion of marine habitats to be found anywhere in Micronesia. 

The Japanese recognized the unusual scientific opportunities pre- 
sented by the Palaus and, during their administration of the South 
Seas Islands (Nanyo Gunto) , established at Koror a tropical biological 
station devoted to the study of coral reefs, under the directorship of 
Dr. Shinkishi Hatai. At that time, Koror was the capital of the 
Japanese mandated islands of the South Seas and was a thriving city. 
The Palao Tropical Biological Station is no more, but we hope to build 
upon its accomplishments and contribute further to a more thorough 
understanding of the fauna of Palau. 



ACTIVITIES IN THE FIELD 

Our first big job on Koror was to unpack and organize our field 
equipment. The territorial entomologist, Kobert P. Owen, had gen- 
erously provided us with laboratory space in the entomological labora- 
tory (pi. 2, fig. 2), one of the few bullet-scarred Japanese buildings 
still usable. A rebuilt second story over the laboratory served as our 
quarters until a nearby house could be made ready for occupancy. For 
days we moved boxes and steel drums full of supplies from the district 
warehouse to the laboratory grounds, and the unpacking went on far 
into our first few nights in the islands. 



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PALAU — BAYER AND HARRY-ROFEN 489 

One morning while we were busy setting up the laboratory with 
the help of Bob Owen's Palauan assistants, we became aware of a 
scowling stranger who at first hesitated around the periphery of ac- 
tivity, and then picked up a hammer and began energetically opening 
boxes. This was Rikrik (pi. 1 8, fig. 1 ). We asked the others about 
him and learned that he was a willing and able worker, knowledgeable 
in English, Palauan geography, and fishing. We had learned from 
experience that such a man is indispensable to an expedition, so we 
hired him on the spot. Rikrik's scowl broke into a broad but tempo- 
rary grin and we had gained a true friend. Later, we added to our 
staff another Palauan named Sumang, who had a remarkable knowl- 



edge of Palauan natural history. He could speak both English and 
Japanese, was a village chief or "Ya'at," and knew practically every- 
body from Angaur to Kayangel. Amiable Sumang Y. was a valuable 
public-relations department whose good offices were a great advantage, 
particularly during the long overland trips on Babelthuap, and his 
memory of Palauan geographic names gave valuable documentation 
for our collection records. 

The Palauan people use a different approach from ours to naming 
the various parts of their homeland. They often do not give names 
to islands as a whole, whereas groups of islands or localized regions on 
islands may have special names. Rivers and streams may have as 
many as three names — one for the part near the mouth, another for the 
headwaters, and a third for the parts between. The imposition of our 
own practice of giving a single name to geographical features upon 
the Palauan system has led to either a part taking the name of the 
whole, or the whole taking the name of one of its parts. Examples of 
the latter kind are Koror, which is the name of a village that we apply 
to an entire island, and Eil Malk, the name of a cape which we use 
for the island of which it is a part. The situation is complicated by 
the circumstance that we take many of our spellings of Palauan place 
names from Japanese maps, which expressed them in phonetic 
katakana characters. The English transliterations from the Japanese 
spellings usually bear little, if any, resemblance to actual pronuncia- 
tion, but they appear almost universally on American maps so we are 
obliged to employ them in this account. Thus, the name "Ankosu" 
as we use it is correctly spelled "Nghus," and "Geruherugairu" should 
be "Ngaregelngael." A complete list of the place names we will men- 



tion in these pages, giving the correct (and any common alternate) 
spelling, may be found on p. 507. 

Actual fieldwork could not be started until our 18-foot fiberglass 
boat was put into commission. Sterling H. Pierce, our engineer and 
electronics technician, installed wiring, instruments, and cabin con- 
trols for the powerful outboard motor. In due time, the final coat 



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490 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

of paint was applied, the name Lenore and the George Vanderbilt 
Foundation insigne inscribed on her bows, and the boat was ready 
for launching. She was both speedy and seaworthy, and in her we 
could make trips to outlying islands and return to the laboratory the 
same day, a distinct advantage when perishable specimens must be 
promptly preserved. 

When the Lenore was fully loaded for a day's work, there was 
scarcely room left for us. Nets, containers large and small, inflatable 
floats for receiving specimens as we took them from the water, spray 
apparatus for distributing the poison we used to stupefy fishes and 
other active specimens, tools for digging and breaking up coral, pre- 
servative, cameras and, often, bulky diving apparatus loaded the boat 
to capacity. 



The self-contained diving apparatus was useful not only for col- 
lecting in water too deep for free diving, but also in shallow-water 
areas where we wanted to see the exact situation under which certain 
animals were living, or to observe their behavior at length. When- 
ever possible, both the habitat and the inhabitants at collecting sta- 
tions were photographed in detail, both in color and in black-and- 
white, using reflex cameras in waterproof casings (pi. 7, fig. 2) . These 
excellent cameras greatly minimized waste of film by enabling us to 
watch moving specimens until they were in range and focus. 

Because one of our chief aims was to get as complete a biological 
sample as possible, every available means of collecting specimens was 
employed, from hand capture and hook-and-line to explosive charges. 
Although different situations required different techniques of sam- 
pling that had to be carefully decided upon before attempting to col- 
lect, the most generally useful method of obtaining active specimens 
was by the use of the vegetable poison rotenone. In liquid form this 
is extremely potent, so it must be diluted with water in a spray pump 
and distributed over the area to be collected. Fishes, crustaceans, 
cephalopods, and certain other types of animals are suffocated by it 
and are soon made helpless. When we used this technique, we needed 
every available hand to collect specimens before they were swept away 
by the current or eaten by larger fishes not affected by the poison. 
Often we enlisted the aid of youthful spectators, who are character- 
istically good collectors, and they would scurry about in response to 
Sumang Y.'s commands, enjoying all the bustle of excitement, 

IWAYAMA BAY 



Our program of faunal sampling took us the length of Palau, from 
Arekolong Peninsula at the northern end of Babelthuap to Peleliu, 
but one of the most fascinating and complex areas in the islands was 
virtually in our own front yard. The Japanese scientists of the 



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PALAU — BAYER AND HARRY-ROFEN 491 

Palao Tropical Biological Station were attracted by it 20 years ago 
and made a survey of it that was reported in the first volume of their 
journal (Abe, 1937) . We became greatly interested in that study and 
had decided, even before reaching Palau, to resurvey the same area to 
see what had happened to the different habitats since they were origi- 
nally studied. This area is an island-studded lagoon, partly enclosed 
by Koror Island (pi. 4), which the Japanese called Iwayama-wan, or 
"Eocky -mountain Bay." The Japanese name is generally accepted 
now, because the nearly forgotten original Palauan name for it is a 
matter of debate (some maps use Kramer's coined name for it, "Songel 
a Use," which, although utilizing Palauan words, is a European inven- 
tion). We therefore use Iwayama Bay as a neo-Palauan name. 

Iwayama Bay is a roughly circular body of water about V-/ 2 nautical 
miles in diameter, enclosed by Koror on the north and Auluptagel on 
the south (see map, fig. 3). Its west entrance is a long, narrow pass 
called Lebugol Channel, and its east entrance a wider, coral- and sand- 
choked passage called Ngasaksao Pass. The western arm of Koror 



is volcanic land with a wide, muddy mangrove shore ; its eastern arm 
is limestone, like Auluptagel and the 40 small islands in the Bay, with 
a fringe of corals. The Palauan names for most of the islands are all 
but forgotten (Sumang Y. succeeded in tracking down most of them 
by lengthy conferences with the prominent patriarchs of Koror) so, 
to simplify matters, the Japanese students assigned each island a 
number, which they actually emblazoned on them in white paint. 
These roman numerals are still legible on some of the islands, and we 
also found that system more convenient to use than names, either 
Palauan or Japanese. N. Abe and his colleagues further divided the 
Bay into "divisions" bearing letter designations (see fig. 3), in most 
of which they studied transects from island shore to reef margin. In 
the process of collecting, we revisited each of the transects in the bay 
and studied the 16 divisions, wherever possible taking photographs 
of the coral growth, animal communities, and general habitat. 

Because of the many islands and narrow passes, tidal currents are 
swift at many points in Iwayama Bay. The Islands, which are very 
close together, rise up almost vertically from the bay floor and create 
very narrow, deep waterways many of which are 100 feet or more in 
depth. The islands are deeply undercut at the high-tide line (pi. 6, 
fig. 2) — as much as 5 to 1 feet — forming deep "notches" above a sub- 
marine shelf of variable width. In favorable localities where the cur- 
rent is strong, as on the north shore of island 29, coral growth on the 
shelf and vertical submarine cliff is exceptionally luxuriant. Here, 
one could stand among flourishing corals and look either directly 
overhead into dense jungle vegetation or straight down the coral preci- 
pice into a hundred feet of deep blue water. Occasionally we saw the 



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ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 



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PALATJ — BAYER AND HARRY-ROFEN 493 

shadowy shapes of giant blue parrotfishes, perhaps 6 feet long, rise 
from the depth to which they had fled at our coming, or a shark patrol- 
ling his accustomed beat, or the faint silvery glint of sunlight on a 
distant school of swift predators, perhaps some kind of jack (Car- 
angidae) , near the limit of our vision. 

On the submarine shelf of these undercut shorelines, the coral 
variety was great, with multicolored Pectinia lactuca, commonly called 
lettuce coral (pi. 9, fig. 1), green, orange, or brick-red Lobopkyllia, 
and a few others dominating the shallower areas. The less colorful 
Plerogyra, with bubblelike tentacles an inch across, the closely re- 
lated Physogyra, and long, wiry antipatharians (black corals) were 
usually found on the vertical face of the cliffs. On the shadiest slopes 
we often met with handsome specimens of Palauphyllia, a subgenus 
of corals named in honor of their homeland. 



The bright colors of Palauphyllia, like those of most other stony- 
reef corals, are located not in the coral skeleton but in the soft tissues 
of the polyps themselves, and are due in part to the presence of minute 
unicellular algae living in the cells of the endoderm. These remark- 
able algae, called zooxanthellae, actually serve the coral polyps in the 
capacity of excretory organs by taking up from the animal tissues such 
waste products as they can use in their own life processes. They have 
never been found living free of corals, and have never been artificially 
cultured. Their only reproductive process seems to be simple division 
and they are passed on from generation to generation of corals 
through the eggs, which become infected before leaving the parent. 

Several species of fishes were found only in such situations in Iwa- 
yama Bay. Among these were the brilliant gold-and-black striped 
butterflyfish (Chaetodon octofasciatus Bleeker), wrasses of the genus 
Gheilinus (pi. 10, fig. 2) , and several kinds of cardinal fishes belonging 
to the genus Apogon. Many species of highly colored fishes that reach 
a length of no more than one or two inches balance like jeweled 
spangles among coral branches that rival them in beauty, or conceal 
their splendor in holes in the coral cliffs. Ever present are the preda- 
tors that seek out these defenseless inhabitants of the coral slopes. 
Some, like trumpetfish, have long snouts with which they relentlessly 
explore all holes and crevices in search of prey; others, such as the 
turkeyfish, have wide jaws and gaping mouth that enable them to en- 
gulf their prey in one quick gulp, in much the same way that a 
vacuum-cleaner may inhale a feather. Solitary sharks were patrolling 
the bottom of the deep waterways, but their presence was of most 



importance to the inhabitants of the deeper waters for the sharks sel- 
dom came to the surface. 

Some of the islands in Iwayama Bay have protected little bay lets 
in which the coral growth may consist largely of Goniopora, a thick- 



[Begin Page: Page 494] 

494 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

branched stony coral with huge, protruding polyps (pi. 10, fig. 1). 
We soon learned to expect trouble when we collected these secluded 
spots, for the unusually virulent nematocysts, or stinging capsules, of 
the Gordopora polyps were easily dislodged by the currents we pro- 
duced by swimming, and stung us so badly that we sometimes were 
obliged to leave the water. 

Much more troublesome than Goniopora, however, was an arbor- 
escent, colonial scyphistoma (called Stephanoscyphus) of the medusa 
Nausithoe, which also released its nematocysts into the water upon 
agitation by strong currents. The stings produced by this animal left 
angry red welts that itched for days afterward, and even caused swell- 
ing of adjacent lymphatic glands. It seemed probable to us that a 
dense growth of Stephanoscyphus could liberate enough nematocysts 
into the water to inflict serious stings upon unsuspecting swimmers. 
Fortunately, this coelenterate seems to be common only locally in 
Palau, where we found it in only two localities. 



In spite of seemingly ideal surroundings in most parts of Iwayama 
Bay, dead corals could be found at almost any location, suggesting 
that conditions are not always so favorable. A day or so of heavy 
rainfall will dump tons of fresh water not only into the bay but also 
upon the islands, from which it cascades down to the Bay, carrying 
with it great quantities of silt and forest debris. Salinity must be 
much reduced, especially near the surface, for many hours, if not 
days; the normal water temperature of about 85° F. may be lowered 
by 5° or more ; suspended matter beclouds the water and the more deli- 
cate corals may be smothered. But, after a few clear days and several 
tidal cycles, the water clears and the survivors continue their struggle 
for existence. 

The northwest corner of the Bay, forming divisions M, N, and O, 
is bathed by good tidal flow, but silt from the nearby mangrove 
shore discourages coral growth and the reef has a sickly appearance 
that belies the large number of species that comprise it. One of the 
resurveyed transects crosses the reef-flat at the south end of island 15, 
and we studied it during several low tides. Here we collected dead 
coral heads with cylindrical black sponges growing on them, which 
we soon found were only the external portions of a boring sponge 
that had excavated great hollow caverns in the coral boulders. Some 
coral heads were thus reduced to hollow shells, and we have yet to 
learn what becomes of the sponge after it has completely "eaten itself 
out of house and home" — whether it then assumes a massive, free- 
living form as do some other boring sponges, or simply dies of expo- 
sure. Whatever its fate after the destruction of the rock in which 
it lives, it is certainly an active reef -destroying agent. At the same 



locality we found another rock-boring sponge that attacked not only 



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PALAU — BAYER AND BARRY-ROFEN 



495 



dead corals but also the limestone floor of the tide-level notches. 
This sponge is almost certain to prove to be one of the important bio- 
logical agents contributing to the formation of undercut shorelines. 
The swift tidal currents that breath life and variety into the reefs 
do not reach the upper part of division K in the northeast corner of 
Iwayama Bay, which is a quiet backwater. An abundant growth of 
coral is nevertheless present, but it lacks variety. Fingery masses 
of Porites and great, white, papery chalices of Montipora (pi. 1 1 , 
fig. 2) flourish everywhere, but few others can be seen. Even the 
fishes are fewer in kind and smaller in size. Small dragonets (Cal- 
lionymidae) dart about in sandy patches or seek refuge among the 
coral branches. One of these dragonets {Synchiropus splendidus) is 



Figure 4. — The splendid dragonet, Synchiropus splendidus, slightly larger than natural size. 

probably the gaudiest fish we found during the summer — colored blue, 
green, and red in an intricate design of spots and bands (see fig. 4). 
A few species of damselfishes (Abudefduf) were abundant, each fish 



with its own territory of coral and water. Each individual vigorously 
defends its own particular home, bullying intruders with threatening 
advances, cheeks distended, clicking and grunting indignantlv. 
Hovering over the branches of Pontes we could always find little 
bronze and maroon cardinalfishes {Apogon nematopterus) , ready to 
retire into their stony sanctuaries at the approach of our staring 
camera lenses (pi. 1 1 , fig. 1 ) . 

In the undercut shoreline of Koror nearby we found a small cave 
of the sort that permeates many of the limestone islands (pi. 6, fig. 1 ) 



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496 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

It was too small to crawl into very far, but in its dark and narrow 
recesses we found some alcyonarian corals (Telesto) hanging from 
the roof, completely exposed by the ebbing tide. Elsewhere there are 
larges caves, some at water level and large enough to admit a boat, 
others equally large but completely submerged. In most of the major 
limestone islands there are large caves that have collapsed, forming 
lakes connected with the sea by subterranean passages. These salt- 
water lakes contain large gobies, mussels, and sedentary jellyfish 
(Cassiopeia). 

One of our reconnaissance techniques was to cruise slowly along 
the coral slope in the Lenore, watching for changes in the appearance 



of the reefs to indicate interesting spots to be examined more closely. 
We followed the bay shore of Koror south from the little cave, around 
the so-called Arappu Peninsula (Ngalap), where the submarine cliff 
was almost devoid of corals and the only conspicuous organism was a 
large, sprawling, branched, pale pink sponge that looked white in 
20 or 30 feet of blue water. Even the stiff snaky "wire corals" 
(Antipatharia) that usually thrive on the cliffs were missing. We 
could find no explanation for the absence of corals there, for the 
current flow is better than it is in many other places, the water clearer. 

After we passed through the narrow strait called Kaki-suido 
(Oyster Pass) in the Japanese reports (Palauan name Ngerikiuul), 
the situation changed, and we came upon one of the most interesting 
areas in Iwayama Bay, one that we visited and revisited, each time 
to find something new. Here the undercut was very deep and the 
foliage of the jungle-covered slopes above hung far down over the 
water, blocking the midday sunlight and producing an almost con- 
stant twilight. For only a few moments in the afternoon could a 
few rays of sunshine slip through before the shadow of island 29 
across the pass crept up to throw the waters into increasing darkness. 
A yawning cavern gaped in the cliff wall, entirely under water, its 
roof festooned with huge, netlike antipatharians ("black corals") 
hanging down like drapes. They were so flexible and so large that 
they could grow only in hanging position (pi. 18, fig. 2). When 
fresh, their polyps were brilliant orange in daylight, but they ap- 
peared almost white in the murky blue water of the cave. 

ANIMAL PARTNERSHIPS 



Only on this half-lit slope did we find the whip-corals, Junceella; 
they grew here like tall, waving grass reaching out toward the light, 
with their tips drooping like buggy whips. They form the hub of an 
interesting association that we will describe in detail in a future paper. 
The most unusual member of this association is a little transparent 
pink goby, a new species allied to the genus Cottogobius. It is a tiny 



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PALAU — BAYER AND HARRY-ROFEN 497 

fish, not much more than an inch in length, that clings to the stalks 
of the whip-corals. They may be pursued up one side of the coral 
and down the other, but rarely will they move to another coral even 
though it is very near. Our ingenious Rikrik found that by sliding a 
coral stalk through his fist, he could catch its fish in a small net as it 
popped off the end. The water was often murky and the light always 
poor, but we did get a picture of Rikrik catching gobies in a field of 
junceellas (pi. 16, fig. 1) . Also clinging to the whip-corals were many 
handsome gray feather-stars, or stalkless crinoids (comatulids) , some 
of them grasping several corals at once (pi. 1 6, fig. 2). The cirri (the 
clawlike "feet") of the crinoids irritated the surface of the corals and 
scarred it permanently wherever they had clung, indicating that these 
"free-living" crinoids move about little, if at all. One of the crinoid's 
distant cousins, a bright orange brittle-star, lived entwined in the arms 
of the crinoid, and among them could also be found a spider crab, 



Harrovia, that had joined the partnership. Another crustacean, a 
small porcellanid crab, scuttled over the surface of the whip-corals, 
completing this curious association of vertebrates and invertebrates. 

Among the most studied of symbiotic relationships to be found on 
tropical reefs is that involving several species of large sea-anemones 
that allow certain kinds of small but colorful damselfishes (genera 
Amphiprion and Dascyllus) to seek protection among their tentacles. 
These clownfishes, as they are sometimes called, rarely stray far from 
their host anemone, and are ready to dart down among the stinging 
tentacles at the first hint of danger (pi. 14, fig. 2). In spite of years 
of study, the details of this association are still not clear (Gohar, 1 948 ; 
Gudger, 1946). It is believed that the fishes avoid being stung by 
swimming in a distinctive fashion that is "recognized" by the coelen- 
terate. Clownfishes have been seen to drag food to the waiting ten- 
tacles of the anemones, but, on the other hand, we watched an Amphi- 
prion seize a tentacle of its host in its mouth and with a few quick 
tugs pull it loose and eat it. This finny ingrate expected, and received, 
sanctuary from the very anemone it had been nibbling upon, for it 
dashed headlong among the tentacles when we approached too closely. 
Some investigators have suggested that by eating bits of the anemone, 
Amphiprion builds up an immunity to nematocyst poison, but this 
suggestion has never been scientifically confirmed. It does seem fairly 
certain, however, that the clownfishes recognize their preferred species 
of host anemone partly by sight and partly by chemical emanations. 
There is also some indication that the anemones do not sting their 
partner fishes because of some kind of chemical "recognition." 

Coelenterates and echinoderms seem often to play the host role in 



these partnerships, probably not because of any inherent good nature 
or native generosity, but because they are slow moving or sedentary 



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498 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

and therefore an easy mark. One of our most startling discoveries 
of the summer was a fish that lives with an echinoderm. This, of itself, 
is not unusual, for the eel-like pearlfishes (Carapus, Fierasfer, and 
Jordanicus) have long been known to inhabit the cloaca of holo- 
thurians (sea-cucumbers) and the body cavity of Culcita (the cushion 
starfish) . But our fish was a previously unknown species of clingfish 
(Gobiesocidae) and the echinoderm host was a feather-star (Coman- 
thus), a completely unexpected combination. The clingfish was black 
with a bright yellow stripe down each side, perfectly camouflaged 
among the arms of its host. 

It was on August 7 that we discovered it, as we were returning from 
a 2-day trip to Ngemelis. We had stopped in Meharehar, the labyrin- 
thine lagoon of Eil Malk, to look for future collecting sites and to ob- 
tain some samples of the lagoon bottom sediments. It was a stormy 
day with heavy downpours that had hampered our observations and 
dampened the spirits of everyone aboard the Lenore. We had taken 
the bottom samples in the rain, and were heading for home by way of 
the inside route west of Urukthapel when we found ourselves over some 
coral flats near Ankosu Point, the southernmost cape of Urukthapel. 



Not wishing to pass up any likely localities, we dropped anchor and 
went over the side to look around. The water was about 6 feet deep 
and the bottom was covered with a tangle of staghorn coral 
(Acropora), most of it lying loose upon the sandy bottom. A few 
knolls of massive coral could be seen, with chalice-shaped acroporas 
and sea-fans (Melithaea) growing on them. Here and there, hidden 
among the corals, we found a many-rayed spiny starfish (Acanthaster 
planci), which is a real danger to bare feet (even those tough enough 
to disregard the jagged coral). It is widely feared by the natives of 
Micronesia, and with ample justification. A friend of ours was vir- 
tually incapacitated for a week or more by wounds inflicted by this 
animal, and he was not fully recovered for a month or more. 

Between the coral branches everywhere, and on the coral knolls, 
the restless, fernlike arms of feather-stars swayed with the rising 
tide. Because we knew that crinoids are ever-gracious hosts to a va- 
riety of invertebrates, we collected some of them to find their lodgers. 
There are usually two kinds of shrimp, a galatheid (or "squat-lob- 
ster"), and a polychaete worm, all protectively colored to match their 
host — usually black and greenish yellow in this locality. The first 
feather-star that Adair Fehlmann collected had some black-and- 
yellow striped shrimps among its arms. Safely inside a glass vial, 
they gave us a real surprise. They were not shrimps at all, but 
fishes — and clingfishes at that — the only ones we would find all sum- 
mer. A careful search disclosed a number of additional specimens 
before the current became so swift that we could work no longer. We 



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PALAU — BAYER AND HARRT-ROFEN 499 

continued our homeward trip with one of the prize catches of the 
expedition — some tiny black-and-yellow fishes little more than an inch 
in length. 

In practically any protected sandy area of the lagoon we could ex- 
pect to find the unusual partnership of shrimps and fishes that we 
first observed at If aluk Atoll in 1953. We were delighted to find this 
association at Palau in such shallow water and in such abundance 
that we could observe it closely and collect the animals in numbers. 
At Palau, two kinds of alpheid shrimps and at least four kinds of 
gobies live together with identical habits. 

Each pair of shrimps excavates its own burrow and then plays 
host to a pair of fishes. The fishes sit just outside the mouth of the 
shrimp burrow (pi. 14, fig. 1) while the shrimps repair and deepen 
it, bringing load after load of sand to the surface on their large claws, 
bulldozer-fashion. But should any danger threaten, the gobies dart 
down the hole in a trice, tumbling the shrimps over in their haste. 
From this activity the shrimps detect that something is amiss and 
cease digging until the gobies regain their composure and their usual 
position on the front doorstep. Whether this inadvertent warning 
is the only benefit derived by the shrimps from the association we are 
unable to say at present. It seems probable that the gobies obtain, 
in addition to shelter, an occasional banquet at the expense of their 



hosts, since one of the specimens collected was stuffed with larval 
crustaceans — probably the young of the shrimps whose home it had 
shared. 

There is still much to be learned about these and various other rarely 
observed biological associations, and it will take patient observation, 
study, and experimentation in the field before we know the exact 
nature of the relationship between the partners and how it may have 
developed. 

THE OUTER REEFS 

The outer reefs have an entirely different appearance from the 
lagoon reefs of Iwayama Bay and the staghorn coral thickets of the 
shallow passes. On the west side of the archipelago the reefs are 
barriers, but on the east they are fringing reefs that follow the land 
closely, with an offshore barrier in only a few places. Whether barrier 
or fringing reefs, they are bathed with the always clean water of the 
open sea and pounded by its sometimes thunderous surf. Different 
and stronger corals live in these exposed situations, and a whole new 
population of fishes swims among their branches. The surgeonfishes, 
butterflyfishes, and wrasses that live here are for the most part 
peculiar to this zone of churning, turbulent water. Few species of 
the quiet lagoon waters are hardy enough to adapt to this rigorous 
environment. Sharks and barracudas in particular prefer this region, 



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500 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

constantly patrolling it for food. Certain surgeonfishes also are char- 
acteristic of the surf zone, among them Acanthurus lineatus, a yellow 
and neon-blue striped fish that loves turbulent, surging water. Their 
relatives, the fan-finned zebrasomas, hide under the spreading table 
corals. Huge parrotfishes, six or more feet in length, graze like 
cattle upon the corals, producing a noisy chorus as they crunch their 
limy meals in large, gregarious schools. Moray eels (Gynmothorax) 
lie in wait in their holes, their malicious eyes alert for any unwary 
fish that may pass within striking distance. The butterflyfishes 
(Ghaetodon), so named because of their brilliant coloration, are 
especially numerous and are endowed with an insatiable curiosity. 
They seem unafraid of divers and will approach very close in order 
to get a good look. On many occasions they have startled us by a 
quick nip or a tug on some especially attractive hair. This has an 
electrifying effect upon swimmers already fearful of a surprise attack 
by prowling sharks, and the sheepish feeling combined with relief 
upon seeing these small challengers, is more amusing now than it 
was on the reef. 

The spiny lobsters (Panulirus) that lurk in crevices of the outer reef 
are different species from those found in the lagoon. So are the coral 
crabs (Trapezia) among the massive branches of pocilloporas, veri- 
table giants by comparison with their relatives in more protected 
waters. 



Any specimens taken from the surf zone are collected only through 
great exertion, for it is one of the most difficult habitats to sample. 
Swift currents rapidly dissipate the rotenone and sweep away the speci- 
mens killed by it. The collector himself may be picked up by the 
swell and dropped 1 5 or 20 feet away, as likely as not on a jagged 
coral. "When explosives are used instead of rotenone, all the nearby 
sharks materialize as if from nowhere, considering its sound a dinner 
bell inviting them to a free meal. Needless to say, we used this explo- 
sive technique infrequently, so as to avoid becoming the piece de 
resistance of some shark's luncheon. 

Near the reef passes and in the deeper waters offshore the corals 
flourish in stony gardens of eerie and awesome beauty. Some species 
of Acropora produce towering spires and antlers (pi. 7, fig. 1) that 
bring to mind the reefs made by the same genus of corals in the West 
Indies, and others form great, spreading disks and platforms com- 
posed of tiny branchlets, a growth form peculiar to the Indo-Pacific 
area (pi. 13). The sea gardens of swaying alcyonarians (sea-fans 
and sea- whips) that give the West Indian reefs their color and fluid 
beauty are nowhere to be seen, for the alcyonarians here are nearly all 
massive, rubbery kinds (pi. 13, fig. 2) that have not the elegance of 
their Caribbean relatives. The few species of sea-fans that do occur 



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PALATJ — BAYER AND HARRY-ROFEN 501 



are more abundant along deep channels or in the protected lagoons 
than on the seaward reefs. 

EEL-GRASS ENVIRONMENT 

A conspicuous feature of the Palau lagoon is the great extent of 
sandy bottom covered with eel-grass {Enhalus acoroides and some re- 
lated species) . It is a distinctive and complex habitat. The most ex- 
pansive eel-grass beds lie near Peleliu, to the south of Koror, al- 
though the west coast of Babelthuap also has some fine ones. We 
made two trips to Peleliu to survey the grass beds there, but on both 
occasions we found poor conditions due to the stage of the tide. The 
water was so murky and full of plant debris that collecting was un- 
pleasant and photography impossible. 

Among the animal inhabitants, fishes are particularly abundant and 
thrive in the eel-grass environment. Sharks, jacks, barracudas, and 
other predaceous fishes constantly search the eel-grass beds for prey. 
Food is not easy to find here, even in the midst of plenty, for the eel- 
grass forms the home of many highly specialized fishes that blend with 
their background in both form and color. Some of them, such as the 
pipefishes (Syngnathidae) and certain wrasses (Cheilio), are elongate 
in outline and green in color, so perfectly camouflaged that sharp eyes 
are needed to separate them from the grass in which they live. Others, 
such as the parrotfish (/Scarlchthys), spinefish (Siganus), and some 
snappers (Lethrinus), are not shaped like the grass blades but are so 
much like it in color that they are virtually invisible. The filefishes 
(Monacanthus) , blennies (Petroscirtes) , and dragonets {C allionymus) 



go a step farther in having their bodies covered with waving filaments 
and hairlike growths that resemble the hydroids and other epiphytes 
covering the eel-grass blades. 

The Palauan boys pointed out some peculiar little black-and-yellow 
fishes that were swimming about a water-logged branch half buried 
in the sand and hidden by grass and warned us that they were very 
dangerous. When we insisted upon catching them, Sumang Y. and 
Eikrik must have regretted pointing them out to us. The fish were 
small catfish (Plotosus anguillaris) with barbed pectoral and dorsal 
spines that are venomous and can inflict a nasty wound. They were 
swimming in a curious manner, very close together and wiggling vigor- 
ously, in a compact school that moved slowly forward like a dark 
cloud. It was simple to frighten them into the range of a large dip- 
net, and each fish captured caused our sturdy Palauans to wince in 
anticipation of the painful punctures to come when we pulled them 
out of the nets, and later, when we placed them in containers of for- 
malin. A month or so later, when we were in Japan to consult with 
biologists who had worked in Palau before the war, we encountered this 



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502 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

same fish in southern Honshu. It was living in a completely different 
ecological situation, in rocky tide pools near the Seto Marine Biologi- 
cal Station at Shirahama. 



One of the major eel-grass inhabitants is a mammal, the little-known 
dugong, or sea-cow (Dugong dug on). During our stay in Palau we 
were always especially watchful while passing over eel-grass areas, 
in the hope of seeing some dugongs, but as they are quite scarce we 
never did see one in its natural habitat. During our last few days at 
Koror, however, some Palauans speared and captured alive a half- 
grown specimen that provided us with considerable excitement and be- 
came the first ever to be exhibited alive in an American aquarium. 
We bought this dugong, otherwise destined for sale as food in the local 
market, and kept it alive in a large pool at the end of Koror dock 
until we were ready to begin our return trip to the United States. 
Then we caught it, wrapped it in wet blankets (pi. 20, fig. 1), and 
carried it, lying on the floor of the plane between our seats, to Guam 
(pi. 20, fig. 2). At Guam it was ensconced in a sturdy crate and 
transferred to a commercial airline for shipment to California. The 
California Academy of Sciences had arranged for its transportation 
to the Steinhart Aquarium, where it proved to be an unusually popular 
exhibit. It was certainly the first Palauan dugong ever to fly to 
America, and we have no doubt provided some material for the "talk- 
ing picture" carvings that decorate Palauan abais. 

HERPETOLOGICAL STUDIES 

Another Palauan animal that never before had been displayed alive 
in an American zoo or aquarium, and which we succeeded in bring- 
ing back to the United States, is the deadly poisonous sea snake 
(Laticauda colubrina) , a relative of the cobras and coral snakes. 



Our first encounter with the banded sea snake in the wild was on 
a field trip to Ngemelis, a group of islands along the southwest barrier 
reef. We were making our way toward the beach just before dusk 
when, a hundred yards or so from shore, we came upon a huge snake, 
a good 6 feet in length, slowly working its way seaward along the 
bottom, poking its head into nooks and crannies and in and out of 
corals, carefully feeling with its tongue. It paid no attention to us, 
and we stood or swam near it for several minutes, in water perhaps 
5 feet deep. At no time did we see it surface for air, and as we con- 
tinued on our way it was still swimming seaward along the bottom. 

The Palauans have an odd story about using sea snakes to catch fish. 
They say that if one holds the snake by the tail, it will probe among 
the rocks and catch fish, which can then be easily seized. We never 
saw this method of fishing practiced. Probably no modern Palauans 
are courageous enough to try it, for our helpers invariably let out 



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SMITHSONIAN REPORT, 1956— BAYER AND HARRY-ROFEN 



•m 



PLATE 1 1 

1 . Graceful, spotted cardinalfishes (Apogon nematopterus) hover over the branches of a 
Pontes colony that is their refuge in times of danger. This little fish, with its red spots 



on a bronzy ground color, and a dark-brown cummerbund, is one of the loveliest denizens 
of protected reefs. 



)V- A 



2. Huge, fragile chalices of Montipora foliosa dominate the reef slope in the innermost part 
of K Division of Iwayama Bay. They form an uncertain retreat for fishes like the small 
damselfish (Pomacentrus) swimming above them. 



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SMITHSONIAN REPORT. 1956. — BAYER AND HARRY-ROFEN 

1. The beautiful falls of Matal Eigad on the Ateshi River, in Ngardmau Municipality of 
Babelthuap. Note figure at foot of falls. The only fishes collected above the falls were 
eels, which can travel overland around obstacles insurmountable to other fishes. (Photo- 
graphed by H. A. Fehlmann.) 

2. H. Adair Fehlmann and Rikrik collecting in a small stream on Arekalong Peninsula of 
northern Babelthuap. Such a locality yields many gobies, and prawns of the genus 
Macrobrachium. 



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SMITHSONIAN REPORT. 1956.— BAYER AND HARRY-ROFEN 



1 . Great, circular plates of acroporas grow one above the other like modernistic buildings 
in this scene along a reef channel south of Ngaremediu. 

2. The convoluted mass in the foreground is a massive, fleshy alcyonarian, Sarcophyton, as 
large as a bushel basket. This type of alcyonarian is typical of Pacific reefs, whereas in 

the Atlantic the bushy gorgonians dominate. 



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SMITHSONIAN REPORT, 1956.— BAYER AND HARRY-ROFEN 

1 . At left, a pair of speckled gobies sit outside the burrow made by the dark-banded snapping 
shrimps. At right, the banded gobies are living with a pale, fine-striped shrimp that is 
almost invisible against the sand. 

2, At left, a clownfish (Amphiprion) pauses above its host anemone. At right, a clownfish 
can be seen hiding among the tentacles of the anemone, which do not sting it. 



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SMITHSONIAN REPORT, 1956. — BAYER AND HARRY-ROFEN 



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SMITHSONIAN REPORT, 1956 — BAYER AND HARRY-ROFEN 



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SMITHSONIAN REPORT. 1956. — BAYER AND HARRY-ROFEN 



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SMITHSONIAN REPORT, 1956— BAYER AND HARRY-ROFEN 



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SMITHSONIAN REPORT. 1956. — BAYER AND HARRY-ROFEN 



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PALATJ — BAYER AND HARRY-ROFEN 503 

great yelps of fear whenever we caught sea snakes (or any other kind, 
for that matter), and their repugnance extends also to eels, perhaps 
with more justification. Another droll story, frequently seen on the 
carved "talking picture" boards, tells how the sea snake taught 
Palauans to fish with nets. In this tale, a Palauan woman, whose 
children were a banana tree, a pussycat, and a sea snake, grew weary 
of her offspring and told them to make their own way in the world. 
The banana tree pointed out the impracticality of this suggestion, at 
least from its rooted standpoint, and thus won a reprieve, but the two 
more active "children" were sent packing. As they swam toward 
Orukuizu, the cat riding on the snake's head, they grew hungry, and in 
Palauan carvings we see the snake encircling some fishes so the cat 
could catch them as they swam out through the narrow space between 
the snake's head and tail. A villager who was up in a palm tree 
gathering toddy caught sight of this procedure and at once recognized 
its possibilities. He rushed down to the shore and asked the snake to 



show him how to fish that way, so the snake showed him how to set 
up a fence of sticks in shallow water, in which fish would be trapped. 
But the Orukuizu man still needed the cat to catch the fish as they 
passed through the mouth of the weir, so he asked the snake if he could 
have the cat, too. The snake agreed but made the man promise to give 
the cat one of the fish each time they made a successful catch. Such 
was the origin of cats and fishweirs. 

This particular species of sea snake is not so completely aquatic as 
are some of its relatives, and it could be found on the seaward shore of 
exposed islands, coiled up in underbrush on dry land or even in the 
branches of trees (pi. 1 9, fig. 2) . They were very common at Nardueis, 
off the southeast shore of Babelthuap, where we caught as many as a 
dozen in a couple of hours. Some of these are still living in the Stein- 
hart Aquarium in San Francisco, but the specimens destined for the 
National Zoological Park in Washington perished during the flight 
across the continent. 

The herpetological collections we assembled contain at least four 
other species of snakes, including a burrowing form (Typhlops) no 

The story of the sea snake, the cat, and the banana tree. Adapted from an actual Palaun 



carving. 



412575 — 57 83 



[Begin Page: Page 504] 

504 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

larger than a moderate-sized earthworm. Geckos (nocturnal lizards 
with adhesive toe pads that aid in climbing) of several kinds abound 
in the trees and on window screens of buildings where they catch in- 
sects attracted to electric lights at night. The toad is extremely 
abundant in the vicinity of Koror, and two kinds of frog seem to 
be widespread throughout the islands. The monitor lizard is not 
especially common but may be encountered on Babelthuap. The 
collections and notes of Dr. Masamitsu Oshima, who studied the rep- 
tiles of Palau, were unfortunately destroyed during the course of the 
war before they could be published. Our collections will therefore 
fill an important gap in our knowledge of the terrestrial fauna of the 
Palaus. 

HOMEWARD BOUND 

By the time November rolled around and Project Coral Fish had 
been active in Palau for four months, we began to make plans for 
the homeward journey. On November 4 we made our last marine 
collecting station, and on the following day the dismantling of the 
laboratory began. All equipment had to be either packed for ship- 
ment or stored for use by Project Coral Fish II. Outboard motors 
were cleaned and boxed, the Lenore was stripped and crated ; collec- 
tions were packed in tins, drums, and boxes, labeled, and stacked up 
to await the next freighter to Guam. In a few days the laboratory 



was starkly in order and deserted. Arrangements were made for 
the transportation of the live cargo of sea snakes and dugong. "Plane 
Day" on November 15 was a typical rainy-season day in Koror. Rikrik 
and Sumang Y. went early to the dock and caught the dugong, and 
it was difficult to say if it was wetter in the pool or on the dock. 
Dugong, snakes, other precious collections, and ourselves were loaded 
on the Albatross amphibian in the still-pouring rain, and we were 
off, it seemed after having scarcely arrived, so swiftly did the months 
pass on these coral-fringed islands and broad lagoons. Only yester- 
day, so it seemed, we had made our way up one of Babelthuap's 
rivers as the crocodiles slid off the banks into the dark water, 
or had been listening to the little bird with a "pipe sweet as a 
flageolet" as we collected in some forest-bound coral bay. Perhaps 
nowhere else in the world are reefs and jungles so intimately asso- 
ciated that one can stand among living corals and at the same time 
collect many different kinds of orchids from overhanging branches. 
It will be many a year before the complexities of this multifarious 
habitat are thoroughly understood, but we think that Project Coral 
Fish is making a long stride toward that understanding and, there- 
fore, toward a better understanding of life in the seas as a whole. 

During the 1055 expedition to Palau, Project Coral Fish assembled 
thousands of specimens from 276 collecting stations extending from 



[Begin Page: Page 505] 



PALAU — BAYER AND HARRY-ROFEN 505 

the northern end of Babelthuap to Peleliu. Phylogenetically, the 
collections contain animals ranging from the Protozoa to the verte- 
brates, as well as many specimens of marine plants. The largest 
collections are those of fishes, crustaceans, and sponges, but the smaller 
collections are no less significant. Taken together, they represent a 
contribution toward a more complete faunistic and zoogeographic 
knowledge of the Palaus, an especially significant one in view of the 
fact that the collections made in the same territory by Japanese 
scholars prior to World War II are either widely scattered over Japan 
or were destroyed during the war. In addition to their systematic and 
zoogeographic value, these collections include information on reef 
communities and biological associations, which forms the framework 
of reef ecology. In spite of its encouraging success, the first field 
season of Project Coral Fish did not accomplish all there was to be 
done in Palau. Neither will the 1 956 season. Some of the smaller 
component projects will certainly see early completion, but they bring 
into bold relief many other problems, not a few of which demand 
an experimental approach that can succeed only at a place like Palau. 

CHARACTERISTICS OF THE PALAUAN MARINE FAUNA 

The large collection of marine animals assembled in Palau during 
the summer and fall of 1955 is only now being studied in detail, 
so it is not yet possible to analyze the fauna from a zoogeographic 
standpoint. However, some impressions of both its richness and its 
affinities were inescapable during the course of observing, collecting, 
and preserving the many specimens handled. These impressions, 



superimposed upon what we already know from the literature, enable 
us to reaffirm the East Indian relationships of the Palauan marine 
fauna. 

Many East Indian fish groups not common in the oceanic islands 
were found to be abundant in the Palaus. Especially noticeable is 
the archerfish (Toxotes), which travels in large schools along the 
lagoon shores, particularly in mangrove regions, where it penetrates 
the narrow waterways and ascends a considerable distance up streams 
and rivers. Glassfishes (Arribassis) and spotted scats (Scatophagus) 
are abundant in the mangrove swamps and lower reaches of streams 
but are not found in the limestone islands of the southern Palaus. 
Fresh-water eels (Anguilla) grow up to 5 feet long in even the 
smallest streams. Some groups of fishes, such as the snappers {Plecto- 
rhynchus, Caesio, Lethrinus, and others) are represented by many more 
species than are known from the multitudes of oceanic islands in the 
Marshalls, Marianas, and Carolines. 

Among the invertebrates, the only two shallow-water gorgonaceans 
(Octocorallia) to extend an appreciable distance eastward into Micro- 



[Begin Page: Page 506] 

506 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

nesia, Subergorgia mollis and Rumphella antipathes, are common 
East Indian species that occur also, as expected, in the Palaus. An- 



other species of /Subergorgia, two magnificent melithaeas, the so-called 
"Plexaura" ftava, a species of Euplexaura, and Junceella fragilis are 
more or less common in Palau, indicative of close ties with the East 
Indies and Philippine faunas. Among the Alcyonacea, Dendro- 
nephthya occurs in abundance and Studeriotes was found in slightly 
deeper water in the lagoon, both genera being well represented in the 
Philippines. These various octocorallian genera are centered mainly 
in the East Indies-Philippines area, but range also northward to 
Japan and southward to Australia, thence eastward to Fiji, Tahiti, 
and Tonga, but reduced in species peripherally. Furthermore, the 
alcyonarian genera typical of Micronesian reefs, such as Sinularia, 
Sarcophyton, Lobophytum, and Sphaerella, are represented in Palau 
by a greater number of species than is usual in the central Pacific 
atolls. 

The molluscan fauna is notably richer in many elements than that 
of the Micronesian atolls to the east, especially in such genera as 
Murex, Spondylus, and Pecten. Since the distribution of Simnia and 
related gastropods is tied to that of the gorgonians on which they live, 
they accordingly do not extend very far to the east, probably not be- 
yond Palau. However, the large egg-cowry (Ovula ovum), 
which lives on soft-corals, enjoys the much wider range of its hosts. 

The fauna of the Palaus will probably not be found to approach 
the richness of that of the Philippines, East Indies, and New Guinea, 
but it certainly far overshadows that of the oceanic atolls. 

PROSPECT 



As we write these words, Project Coral Fish II is continuing the 
program of investigations embarked upon in 1955. It is anticipated 
that the field phase of a survey of the fresh-water fish fauna of Babel- 
thuap, begun in 1955, will be completed successfully during the pres- 
ent season, and the marine survey continues with special attention to 
areas not previously visited. The research program has been broad- 
ened by the addition of an oceanographer to the field party, so that 
more detailed data regarding the aquatic environment may be 
gathered. 

During Coral Fish III in 1957, we hope to complete our ecological 
resurvey of Iwayama Bay, and to make more detailed studies of some 
of the commensal, symbiotic, and parasitic relationships mentioned in 
the foregoing pages. The results of the 1 956 expedition should permit 
much more efficient observations on the physical environment and en- 
hance the value of the Iwayama resurvey. We hope that it will also 



[Begin Page: Page 507] 

PALAU — BAYER AND HARRY-ROFEN 507 

be possible to inaugurate a program of current and plankton studies, 
especially within Iwayama Bay, to determine the reasons for, and to 
provide a better understanding of, the distribution of various corals 
and other benthic invertebrates within the confines of the Bay. 



ACKNOWLEDGMENTS 

The fieldwork described in these pages was carried out under ar- 
rangements with the Pacific Science Board (NAS-NRC) and the 
Office of Naval Research [contract N7onr-291 (57) J. Work in the 
Palaus would have been impossible without the approval and thorough 
cooperation of the Trust Territory of the Pacific Islands. Especially 
we wish to express our appreciation to High Commissioner D. 
H. Nucker, Executive Officer A. M. Hurt, and their staffs in Guam. 
In Palau, District Administrator Donald Heron extended every pos- 
sible courtesy, and his staff, including Francis B. Mahoney, Robert P. 
Owen, and many others, were wholeheartedly cooperative and inter- 
ested in our program. We likewise cannot overlook the Trustees of the 
George Vanderbilt Foundation and the authorities of the Smithsonian 
Institution, who made our participation possible. 

PALAUAN GEOGRAPHIC NAMES 

The names used in this account are those to be found on the charts 
of the U. S. Hydrographic Office and the maps of the U. S. Corps of 
Engineers. The spellings of these names may bear little resemblance 
to their pronunciations. We therefore offer the following table, the 
left-hand column listing the names as taken from published charts and 
used herein, and the right-hand column listing the phonetic spellings 
by Kramer, or supplied to us by Sumang Y. 

Angaur (Island) a Ngeaur 

Ankosu (Point) Nghus 



Arakabesan (Island) Ngarekobasang 



Arekalong (Peninsula) Ngaregolong 



Ateshi (River) [also Addeido] a Did 



Aulong (Island) a Ulong 



Aurapusbekaru (Island) [also Auluptagel] Ulupsagel 



Babelthuap (Island) Babldaop 



EilMalk (Island) a llmalk 



Geruherugairu (Pass) Ngaregelngael 



Kayangel (Atoll) Nggeiangel 



Malakal (Island) Malagal 



Matal Eigad (Falls of a Did R.) Madal a Jegad 



Nardueis (Island) Ngarduais 



Orukuizu (Islands) Ngerkuid 



Palau [also Palao, Pelew, etc.] Pelau or Belau 



Peleliu Peliliou 



Urukthapel Ngurukdapel 



[Begin Page: Page 508] 

508 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1956 

LITERATURE 
Abe, Nobobu. 

1937. Ecological survey of Iwayama Bay, Palao. Palao Trap. Biol. Stat. 
Stud., vol. 1 , No. 2, pp. 21 7-324, 42 figs. 
Abe, Noboru ; Bguchi, M. ; and Hibo, F. 

1937. Preliminary survey of the coral reef of Iwayama Bay, Palao. Palao 

Trop. Biol. Stat. Stud., vol. 1 , No. 1 , pp. 1 7-35, 1 text fig., 2 pis., chart. 
Abe, Tokihabu. 

1 939. A list of the fishes of the Palao Islands. Palao Trop. Biol. Stat. Stud. 

vol. 1, No. 4, pp. 523-583. 
Davenpobt, Demobest. 

1955. Specificity and behavior in symbioses. Quart. Rev. Biol., vol. 30, 



No. 1, pp. 29-46, 7 figs. 
Gohab, H. A. F. 

1948. Commensalism between fish and anemone (with a description of the 

eggs of Amphiprion bicinctus Riippell). Publ. Marine Biol. Stat. 

Ghardaqa (Red Sea), Egypt, vol. 6, pp. 35-44, 4 pis. 
Gudgeb, E. W. 

1946. Pomacentrid fishes symbiotic with giant sea anemones in Indo-Pacific 

waters. Journ. Roy. Asiatic Soc. Bengal, Science, vol. 12, No. 2, 

pp. 53-76, 2 pis. 
Habby, Robebt R. 

1956. "Eugenie" the dugong mermaid. Pacific Discovery, vol. 9, No. 1 , pp. 

21-27, 5 figs. 
Keate, Geobge. 

1 788. An account of the Pelew Islands, situated in the western part of the 
Pacific Ocean. Composed from the journals and communications 
of Captain Henry Wilson, and some of his officers, who, in August 
1783, were there shipwrecked, in the Antelope, a packet belonging to 
the Honourable East India Company. London. 
Kbamer, A. 



1917-1929. Palau. Ergebn. Siidsee Exped. (Thilenius, ed.). II. Ethno- 
graphie, B. Mikronesien, vol. 3. Pt. 1 , 1 91 7 ; Pt. 2,1919; Pt. 3, 1 926 ; 
Pt. 4-5, 1929. 
Kubaby, J. S. 

1873. Die Palau Inseln in der Siidsee. Journ. Mus. Godeffroy, vol. 1, No. 4, 

pp. 1 77-238. 

1895. Ethnographische Beitrage zur Kenntnis des Karolinen Archipels. 

Leiden. 

Nanyo Gunto Bunka Kyokai (South Seas Cultural Association). 

1938. Nanyo Gunto Shashin Cho (Photo album of the South Sea Islands), 

Tokyo.